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PROCEEDINGS 


OF  THE 


California  Academy  of  Sciences 


Vol.  42 


SAN  FRANCISCO 

PUBLISHED  BY  THE  ACADEMY 
1979-1982 


COMMITTEE  ON  PUBLICATIONS 
Tomio  Iwamoto,  Chairman  and  Editor 

Frank  Almeda 

Daphne  F.  Dunn 

William  N.  Eschmeyer 

Frank  H.  Talbot 


(US  ISSN  0068-547X) 

The  California  Academy  of  Sciences 

Golden  Gate  Park 
San  Francisco,  California  94118 


PRINTED  IN  THE  UNITED  STATES  OF  AMERICA 
BY  ALLEN  PRESS,  INC.,  LAWRENCE,  KANSAS 


CONTENTS  OF  VOLUME  42 

Pages 

No.    1.     WILDER,  D.  DEE.  Systematics  of  the  Nearctic  Ptilodexia  Brauer  and  Bergen- 

stamm  (Diptera:  Tachinidae).  Published  March  2,  1979 1-55 

No.  2.  McCosKER,  JOHN  E.  The  snake  eels  (Pisces,  Ophichthidae)  of  the  Hawaiian 
Islands,  with  the  description  of  two  new  species.  Published  March  2, 
1979 57-67 

No.    3.     CHEMSAK,  JOHN  A.,  AND  E.  G.  LINSLEY.  Review  of  the  Rhinotragini  of 

Mexico  (Coleoptera:  Cerambycidae).  Published  March  2,  1979 69-85 

No.  4.  KAVANAUGH,  DAVID  H.  Studies  on  the  Nebriini  (Coleoptera:  Carabidae),  III. 
New  Nearctic  Nebria  species  and  subspecies,  nomenclatural  notes,  and  lec- 
totype  designations.  Published  December  22,  1979 85-133 

No.    5.     IWAMOTO,  TOMIO.  Eastern  Pacific  macrourine  grenadiers  with  seven  branchi- 

ostegal  rays  (Pisces:  Macrouridae).  Published  December  22,  1979 135-179 

No.  6.  FRITZSCHE,  RONALD  A.  Revision  of  the  eastern  Pacific  Syngnathidae  (Pisces: 
Syngnathiformes),  including  both  Recent  and  fossil  forms.  Published  July  2, 
1980 181-227 

No.  7.  BRADBURY,  MARGARET  G.  A  revision  of  the  fish  genus  Ogcocephalus  with 
descriptions  of  new  species  from  the  western  Atlantic  Ocean  (Ogcocephalidae; 
Lophiiformes).  Published  July  2,  1980  229-285 

No.  8.  Poss,  STUART  G.,  AND  WILLIAM  N.  ESCHMEYER.  Xenaploactis ,  a  new  genus 
for  Prosopodasys  asperrimus  Giinther  (Pisces:  Aploactinidae),  with  descrip- 
tions of  two  new  species.  Published  July  2,  1980  287-293 

No.    9.     ROBERTS,  TYSON  R.  Sundasalangidae,  a  new  family  of  minute  freshwater 

salmoniform  fishes  from  Southeast  Asia.  Published  March  5,  1981    295-302 

No.  10.     ALMEDA,  FRANK.  New  and  reconsidered  species  of  Miconia  (Melastomata- 

ceae)  from  Costa  Rica  and  Panama.  Published  March  5,  1981    303-314 

No.  11.  BRIGGS,  THOMAS  S.,  AND  DARRELL  UBICK.  Studies  on  cave  harvestmen  of 
the  central  Sierra  Nevada  with  descriptions  of  new  species  of  Banksula. 
Published  June  24,  1981 1 315-322 

No.  12.  LINDBERG,  DAVID  R.,  AND  JAMES  H.  MCLEAN.  Tropical  eastern  Pacific  lim- 
pets of  the  family  Acmaeidae  (Mollusca,  Archaeogastropoda):  generic  criteria 
and  descriptions  of  six  new  species  from  the  mainland  and  the  Galapagos 
Islands.  Published  June  24,  1981  323-339 

No.  13.     SMALL,  GREGG,  J.  A  review  of  the  bathyal  fish  genus  Antimora  (Moridae: 

Gadiformes).  Published  June  24,  1981    341-348 

No.  14.  FRICKE,  RONALD.  The  kaianus- group  of  the  genus  Callionymus  (Pisces: 
Callionymidae),  with  descriptions  of  six  new  species.  Published  October  26, 
1981  349-377 

No.  15.  ROTH,  BARRY.  Distribution,  reproductive  anatomy,  and  variation  of  Monade- 
nia  troglodytes  Hanna  and  Smith  (Gastropoda:  Pulmonata)  with  the  proposal 
of  a  new  subgenus.  Published  October  26,  1981  379-407 

[iii] 


No.  16.  JONES,  ROBERT  E.  Food  habits  of  smaller  marine  mammals  from  northern  Cali- 
fornia. Published  October  26,  1981  409-433 

No.  17.  KAVANAUGH,  DAVID  H.  Studies  on  the  Nebriini  (Coleoptera:  Carabidae), 
IV.  Four  new  Nebria  taxa  from  western  North  America.  Published  October 
26,  1981  435-442 

No.  18.     GRYGIER,  MARK  J.  Dendrogaster  (Crustacea:  Ascothoracida)  from  California: 

sea-star  parasites  collected  by  the  ALBATROSS.  Published  May  14,  1982 443-454 

No.  19.  ERWIN,  TERRY  L.  Small  terrestrial  ground-beetles  of  Central  America  (Carab- 
idae: Bembidiina  and  Anillina).  Published  May  14,  1982  455-496 

No.  20.  ROBERTS,  TYSON  R.  The  Bornean  gastromyzontine  fish  genera  Gastromyzon 
and  Glaniopsis  (Cypriniformes,  Homalopteridae),  with  descriptions  of  new 
species.  Published  May  14,  1982 497-524 

No.  21.  ZULLO,  VICTOR  A.,  AND  RAJ  B.  GURUSWAMI-NAIDU.  Late  Miocene  balanid 
Cirripedia  from  the  Basal  Wilson  Ranch  beds  ("Merced"  Formation),  Sonoma 
County,  northern  California.  Published  May  14,  1982  525-535 

Index  to  Volume  42  _  537-550 


tiv] 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  1,  pp.  1-55;  113  figs.  March  2,  1979 


SYSTEMATICS  OF  THE  NEARCTIC  PT1LODEXIA  BRAUER  AND 
BERGENSTAMM  (DIPTERA:  TACHINIDAE)* 

By 

D.  Dee  Wilder 

Department  of  Entomology ,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  941 18 


ABSTRACT:  A  revised  classification  of  the  Nearctic  prosenine  genus  Ptilodexia  Brauer  and  Bergcnstamm  (Dip- 
tera:  Tachinidae)  is  presented.  A  total  of  8,000  specimens  and  type  material  for  nearly  all  species  were  studied. 
All  seventeen  previously  described  valid  species  of  Ptilodexia  are  diagnosed  and  illustrated.  Five  new  species, 
P.  sabroskyi,  P.  pacifica,  P.  californica,  P.  westi,  and  P.  maculata  are  described  and  illustrated.  A  key  to  the 
Nearctic  species  is  presented.  The  following  new  synonymies  are  made:  P.  conjuncta  (Wulp)  (=  Rhynchodexia 
simulans  Wulp);  P.  contristans  (Wulp)  (=  K.  punctipennis  Wulp);  P.  carolinensis  Brauer  and  Bergenstamm 
(=  P.  neotibiaKs  West,  P.  minor  West);  P.  halone  (Walker)  (=  P.  hucketti  West);  P.  harpasa  (Walker)  (  = 
P.  leucoptera  West,  Dinera  robusta  Curran);  P.  rufipennis  i  Mai-quart )  (=  Drxia  cerata  Walker,  D.  albifrons 
Walker,  Rhynchodexia  confusa  West,  K.  translucipennis  West,  Rhynchodexia  dubia  Curran);  P.  major  (Bigot) 
(=  Dexiosoma  fumipennis  Bigot,  Rhynchodexia  fraterna  Wulp,  R.  omissa  Wulp);  P.  incerta  West  (=  P.  proximo 
West;  Rhynchodexia  elevata  West). 

The  biology  of  these  parasitic  flies  is  reviewed  and  possible  host-parasite  relationships  are  discussed.  The 
taxonomic  significance  of  numerous  morphological  characters  in  the  genus  and  the  subfamily  is  discussed.  The 
phytogeny  of  Ptilodexia  and  its  nearest  relatives  is  discussed;  six  species  groups  are  separated,  and  an  evolu- 
tionary tree  presented  for  these  groups.  The  contemporary  and  historical  zoogeography  of  the  genus  is  discussed 
as  it  pertains  to  host  and  parasite  distribution.  A  distribution  map  is  presented  for  each  species  treated. 

INTRODUCTION  larvae  of  certain  scarabaeid  beetles.  They  are  of 

Flies  of  the  genus  Ptilodexia  are  large  calyp-  economic  interest  because  they  are  known  par- 

trate  Diptera  belonging  to  the  Tachinidae,  a  fam-  asites  of  such  Pests  as  PhyUophaga  spp.,  Po- 

ily  of  exclusively  parasitic  flies.   Ptilodexia  Pillia  J<*P°"<™   Newman,  and   Macrodactylus 

adults  are  commonly  collected  on  flowers  during  subspinosus  (Fabncius).  The  genus  is  distribut- 

the  summer  months.  The  larvae  parasitize  the  ed  throughout  the  New  World. 

Although  specimens  of  Ptilodexia  are  com- 

mon  in  collections,  few  are  correctly  identified. 

*  Contribution  from  the  Frost  Entomological  Museum,  Sabrosky  and  Arnaud  (1965)  made  no  attempt 

Pennsylvania  State  University.  This  research  was  supported       to  give  synonymies  Or  distributions  in  their  cat- 
by  the  Agricultural  Experiment  Station  Project  No.  2070,  and       alog    listing   of  27    species    names.    The    special 

constitues  Contribution  No.  561 1  from  that  station.  An  earlier      problems  leading  to  such  confusion  in  Ptilodexia 

version  of  this  paper  was  submitted  to  the  Graduate  School,       ^     j}  djfficu,t      jn  associating  t.he  S6X6S,  (2)  a 

Pennsylvania  State  University,  as  a  thesis  in  partial  fulfill-  . 

ment  of  the  requirements  for  the  degree  of  Doctor  of  Philos-       h'gh  degree  of  mtraspecific  variation,  (3)  an  UH- 

ophy.  usually  low  degree  of  interspecific  variation,  and 

[1] 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


(4)  the  lack  of  consistent  traditional  morpholog- 
ical characters. 

The  purpose  of  this  study  is  to  revise  the  clas- 
sification of  Ptilodexia  on  the  basis  of  all  avail- 
able material,  to  analyze  the  life  history  of  its 
species,  and  to  determine  evolutionary  and  zoo- 
geographical  trends  which  may  apply  also  to 
other  members  of  the  family  Tachinidae. 

MATERIALS 

This  study  was  based  on  over  8,000  specimens 
borrowed  from  various  museums.  In  addition, 
type-specimens  for  most  of  the  known  species 
and  numerous  representatives  of  other  genera  in 
the  Prosenini  were  studied. 

Institutions  loaning  material  used  in  this  study 
were  as  follows:  American  Museum  of  Natural 
History  (AMNH),  Arizona  State  University 
(ASUT),  British  Museum  (Natural  History) 
(BMNH),  California  Academy  of  Sciences 
(CASC),  University  of  California  Berkeley 
(CISC),  Canadian  National  Collection  (CNCI), 
Cornell  University  (CUIC),  University  of  Ne- 
braska (DEUN),  Field  Museum  of  Natural  His- 
tory (FMNH),  Florida  State  Collection  (FSCA), 
Iowa  State  University  (ISUI),  Los  Angeles 
County  Museum  of  Natural  History  (LACM), 
Leningrad  Museum  of  Natural  History  (LMNH), 
Museum  of  Comparative  Zoology,  Harvard  Uni- 
versity (MCZC),  Michigan  State  University 
(MSUC),  Ohio  State  University  (OSUC),  Okla- 
homa State  University  (OSEC),  Oregon  State 
University  (OSUO),  Paul  H.  Arnaud,  Jr.,  Col- 
lection (PHAC),  Yale  University  (PMNH),  Pur- 
due University  (PURC),  South  Dakota  State 
University  (SDSU),  University  of  Kansas 
(SEMC),  Staten  Island  Institute  of  Science 
(SIIS),  University  of  Oklahoma  (SMSH),  Texas 
A  &  M  University  (TAMU),  University  of  Ari- 
zona (UAIC),  University  of  Alberta  (UASM), 
University  of  California  Davis  (UCDC),  Uni- 
versity of  California  Riverside  (UCRC),  Univer- 
sity of  Idaho  (UICM),  University  of  Montreal 
(UMIC),  University  of  Michigan  (UMMZ),  Utah 
State  University  (USUC),  National  Museum  of 
Natural  History  (USNM),  Vienna  Museum  of 
Natural  History  (VMNH),  Washington  State 
University  (WSUC). 

METHODS 

The  male  genitalia  of  Ptilodexia  species  are 
partially  obscured  on  dry,  pinned  specimens.  To 


examine  them,  the  posterior  half  of  the  abdomen 
was  removed,  placed  in  a  solution  of  10  percent 
KOH,  and  heated  until  the  structures  were  suf- 
ficiently softened  to  be  dissected  easily.  They 
were  later  rinsed  twice  with  water  and  twice 
with  acetic  acid,  placed  in  glycerine,  and  ex- 
amined. They  were  stored  in  a  microvial  pinned 
beneath  the  insect. 

Illustrations  of  the  genitalia  were  made  using 
an  ocular  grid.  The  postabdomen  was  anchored 
to  a  small  piece  of  soft  wax  on  the  bottom  of  the 
dish  of  glycerine  to  prevent  it  from  drifting 
about. 

Drawings  of  the  heads  were  made  by  project- 
ing photographic  transparencies  of  them  onto 
drawing  paper.  Manipulation  of  the  projector 
provided  images  of  uniform  size.  The  image  was 
then  traced  with  a  hard  pencil;  the  details  were 
filled  in  after  thorough  examination  of  the  spec- 
imen with  a  dissecting  microscope. 

All  measurements  were  made  using  an  ocular 
grid,  calibrated  by  a  stage  micrometer.  Ratios 
were  calculated  from  these  measurements. 

Because  of  the  extreme  intraspecific  variabil- 
ity of  Ptilodexia,  a  description  which  included 
all  variation  would  be  unwieldy  and  would  be 
similar  for  each  species.  To  make  the  descrip- 
tion more  useful  and  manageable  in  size,  only 
one  specimen,  the  holotype,  is  described. 

A  complete  synonymy  and  list  of  citations  are 
given  for  each  species  included  in  this  study. 
New  species  are  thoroughly  described  and  di- 
agnosed. For  previously  described  species,  only 
a  diagnosis  is  presented.  Known  information  on 
each  species  is  summarized  and  notes  regarding 
types  and  nomenclature  are  presented. 

Intraspecific  variation  is  discussed  thoroughly 
for  all  new  species.  With  previously  described 
species,  variation  is  discussed  only  where  it  is 
necessary  for  species  identification.  Complete 
data  from  each  specimen,  including  sex,  locali- 
ty, collector,  depository,  and  other  information, 
have  been  recorded  by  Wilder  (1976)  and  there- 
fore are  not  presented  here.  Wilder  (1976)  also 
gives  complete  redescriptions  for  all  previously 
described  species  of  Ptilodexia. 

The  scope  of  this  revision  has  been  limited  for 
practical  reasons.  Inclusion  of  the  southern 
Mexican  species  would  have  doubled  the  num- 
ber of  species  treated,  and  the  material  available 
for  these  species  is  wholly  inadequate.  All 
Nearctic  species  are  treated  herein. 


WILDER:  NEARCTIC  PTILODEXIA 


ACKNOWLEDGMENTS 

I  extend  my  sincerest  appreciation  to  Dr.  C. 
W.  Sabrosky,  who  initiated  this  study  and  with- 
out whose  help  this  revision  could  not  have  been 
completed.  Dr.  Sabrosky  also  generously  loaned 
types  and  was  always  available  to  answer  ques- 
tions about  Ptilodexia  and  related  tachinids. 

I  thank  Dr.  K.  C.  Kim  for  his  valuable  assis- 
tance during  this  project.  His  enthusiasm  and 
drive  served  as  a  constant  inspiration. 

Persons  and  institutions  loaning  type  material, 
for  which  I  am  grateful,  are:  Dr.  R.  W.  Crosskey 
(BMNH),  who  was  extremely  generous  in  loan- 
ing 56  type-specimens  for  this  project;  Dr.  R. 
Litchtenberg  (VMNH);  Dr.  V.  Richter  (LMNH); 
Dr.  L.  L.  Pechuman  (CUIC)  who  kindly  loaned 
me  the  types  of  14  species  for  an  extended  pe- 
riod of  time;  also  J.  C.  Scott  (MCZC),  Dr.  P. 
Wygodzinsky  (AMNH),  and  Dr.  F.  C.  Thomp- 
son (USNM)  who  helped  in  uncovering  some 
important  syntypes. 

I  am  indebted  to  Dr.  P.  H.  Arnaud,  Jr.,  and 
the  Department  of  Entomology,  California 
Academy  of  Sciences,  for  providing  facilities 
and  valuable  assistance  during  my  time  on  the 
West  Coast.  Thanks  are  also  due  to  Dr.  D.  C. 
Rentz  and  D.  H.  Kavanaugh  for  their  frequent 
assistance  with  many  problems. 

I  finally  thank  my  husband,  George  Zelznak, 
for  his  unending  encouragement  and  optimism 
throughout  this  study. 

BIOLOGY 

The  larvae  of  Ptilodexia  flies  parasitize  and 
kill  their  scarabaeid  larva  hosts.  The  adult  flies, 
however,  feed  on  nectar  and  they  spend  consid- 
erable time  probing  at  flowers,  particularly  com- 
posites. While  feeding  they  become  covered 
with  pollen  and  probably  act  as  pollinators. 
Ptilodexia  conjuncta  and  P.  agilis  adults  have 
been  observed  pollinating  the  flowers  of  dwarf 
mistletoe  (Arceuthobium  cyanocarpum).  Adults 
of  Ptilodexia  have  been  collected  with  pollinia 
attached  to  the  tarsi. 

Adults  are  collected  in  many  environments. 
Members  of  some  species  are  collected  at  the 
seashore,  while  others  have  been  taken  at  alti- 
tudes as  high  as  3,000  m  in  the  Sierra  Nevada 
and  the  Rocky  Mountains.  These  flies  have  been 
taken  at  UV  light,  Malaise,  and  other  flight 
traps.  But  the  most  productive  method  of  col- 


lecting seems  to  be  sweeping  flowers,  especially 
composites  such  as  Baccharis  and  Solidago. 

The  occurrence  of  specimens  of  Ptilodexia  is, 
as  with  most  parasites,  seasonal.  At  times, 
hundreds  of  individuals  of  one,  two,  or  even 
three  species  may  be  collected  simultaneously 
at  one  kind  of  flower.  In  other  years  the  flies  will 
be  rare — perhaps  representing  the  normal  build- 
up and  decline  of  a  parasitic  population.  In  some 
areas,  such  as  Long  Island,  New  York,  and  Riv- 
erside, California,  specimens  of  certain  species 
have  been  collected  on  the  same  dates  every 
year  for  ten  or  more  years.  However,  both  these 
areas  have  been  extensively  surveyed  regularly 
by  specialists  and  may  represent  the  actual  sit- 
uation, which  is  not  seen  in  other  areas  simply 
because  of  poor  sampling. 

There  is  no  information  on  the  mating  habits 
of  Ptilodexia  spp.  Despite  the  numerous  speci- 
mens collected,  few  have  been  pinned  in  copula. 
Males  usually  emerge  before  females,  but  the 
place  and  time  of  mating  is  unknown. 

The  female  carries  hundreds  of  tiny  larvae  in 
her  abdomen  during  larviposition.  It  is  not 
known  if  she  simply  broadcasts  them  or  if  she 
places  them  directly  on  the  soil.  Neither  is  it 
known  if  the  female  is  able  to  locate  areas  of 
host  density,  nor  if  the  larva  has  the  sensory 
capacity  to  find  a  host.  If  the  larvae  are  depos- 
ited in  ajar,  they  will  wander  along  the  sides  for 
two  to  three  days  before  dying.  First-instar  lar- 
vae of  Prosena  siberita  survive  a  week  or  more 
in  the  soil  (Clausen  1927). 

The  larvae  are  presumably  quite  easy  to  rear 
if  the  host  larvae  can  be  kept  alive  under  labo- 
ratory conditions.  During  the  development  of 
the  parasite  larva,  a  defensive  response  of  the 
host  causes  a  respiratory  funnel  to  appear  at  the 
point  of  attachment  to  the  host.  This  is  a  scler- 
otized  funnel-shaped  structure  which  encloses 
the  caudal  end  of  the  larva  and  can  be  seen 
through  the  integument  of  the  host.  The  larva 
feeds  on  the  internal  fluids  and  fat  body  of  the 
host  and  finally  leaves  the  host  to  pupate  in  the 
soil.  By  this  time,  the  larva  has  ingested  nearly 
all  the  contents  of  the  host. 

Based  on  Davis's  (1919)  data,  Ptilodexia  lar- 
vae overwinter  within  the  host.  The  pupal  stage 
is  quite  short,  cold  temperatures  are  not  re- 
quired to  complete  development. 

The  host  scarabaeid  larvae,  or  white  grubs, 
belong  to  the  subfamily  Melolonthinae,  which 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


includes  such  common  and  economically  impor- 
tant genera  as  Phyllophaga,  Popillia,  and  Mac- 
rodactylus.  The  dynastinine  scarabaeids  of  the 
genus  Aphonus  are  also  parasitized  by  these 
flies.  Champlain  and  Knull  (1944),  and  Peterson 
(1948)  implicate  Ptilodexia  canescens  as  a  par- 
asite of  the  cerambycids  Saperda  calcarata  and 
Rhagium  lineatum. 

Most  species  of  Phyllophaga  have  a  two-  or 
three-year  life  cycle.  Popillia  japonica  and  some 
of  the  other  hosts  have  one-year  life  cycles.  The 
life  cycle  of  Ptilodexia  probably  does  not  exceed 
one  year.  Early-instar  white  grubs  are  attacked 
in  the  fall  and  fed  upon  until  the  following  spring 
or  summer  when  the  parasite  pupates.  Adult 
flies  emerge  shortly  thereafter. 

According  to  Davis  (1919),  when  infested  host 
larvae  are  brought  indoors  in  the  fall,  the  Ptilo- 
dexia larvae  will  continue  their  development 
within  the  host,  pupate,  and  emerge  as  adults  in 
the  winter,  without  interruption.  It  is  thus  pos- 
sible that  in  areas  with  long  seasons,  two  or 
more  broods  could  develop.  My  samples  from 
Texas,  southern  California,  and  some  south- 
eastern states  appear  to  support  this.  Two 
broods  per  year  are  possible  only  if  there  is  an 
ample  supply  of  grubs  of  the  proper  stage  feed- 
ing in  the  soil.  In  these  areas  of  bivoltinism,  vari- 
ant populations  in  the  species  concerned  are  ap- 
parently more  common  than  in  areas  where  only 
one  brood  is  possible.  One  can  extrapolate  to 
the  tropical  regions  where  even  more  genera- 
tions per  year  are  possible,  and  the  number  of 
species  and  the  variation  among  species  is  phe- 
nomenal. 

It  is  not  known  if  any  species  of  Ptilodexia  is 
host  specific,  but  some  species  are  known  to 
have  more  than  one  host.  For  example,  P.  car- 
olinensis  can  complete  its  life  cycle  either  in 
Phyllophaga  rugosa  (fide  Davis  1919,  as  Ptilo- 
dexia abdominalis)  or  in  Popillia  japonica.  On 
the  other  hand,  Ptilodexia  maculata  and  P. 
prexaspes  have  restricted  ranges  and  show  little 
intraspecific  variation — perhaps  indicative  of 
host  specificity.  Ptilodexia  harpasa  and  one  of 
its  reported  hosts,  Macrodactylus  subspinosus, 
are  sympatric. 

Many  questions  remain  unanswered  regarding 
the  relationship  of  host  preference  and  specia- 
tion  in  Ptilodexia.  In  some  species,  local  aber- 
rant populations  are  found  that  only  vaguely  re- 
semble the  typical  population.  The  possibility 
exists  that  these  aberrant  populations  have  shift- 


ed to  a  host  significantly  different  from  the  com- 
mon one.  It  is  conceivable  that  such  a  shift  could 
be  an  early  step  in  speciation. 

Many  factors  involving  the  relationship  of 
host  and  parasite  presumably  influence  the  ap- 
pearance of  the  adult  fly.  These  factors  include 
number  of  fly  larvae  per  host,  instar  of  parasi- 
tized host,  rate  of  host  development,  and  rate  of 
parasitoid  development.  Specimens  of  P.  caro- 
linensis  developed  in  Phyllophaga  grubs  differ 
greatly  from  those  developed  in  larvae  of  Popil- 
lia. 

These  factors  contribute  to  the  extreme  intra- 
specific variability  in  Ptilodexia,  perhaps  ulti- 
mately leading  to  speciation  in  the  group.  Care- 
fully controlled  breeding  experiments  are  needed 
to  help  understand  the  effects  on  the  parasites 
of  the  host  and  host  environment. 


TAXONOMIC  CHARACTERS 

Most  of  the  specific  characters  previously 
used  by  Ptilodexia  taxonomists  are  subtle,  dif- 
ficult to  see,  and  unstable,  sometimes  differing 
not  only  between  individuals  but  also  on  each 
side  of  the  same  specimen.  Of  the  characters 
traditionally  used  to  separate  species,  many 
have  been  either  stable  within  the  genus  (e.g., 
"arista  plumose")  or  different  within  a  species 
(e.g.,  color,  wing  venation).  Most  earlier  work- 
ers lacked  sufficient  study  material  to  recognize 
normal  intraspecific  variation.  Certain  charac- 
ters these  early  workers  used  were  good,  but 
they  are  more  useful  used  in  combination  with 
certain  other  characters. 

Sexual  dimorphism  has  caused  problems  in 
the  taxonomy  of  Ptilodexia.  Abdominal  color 
and  color  pattern,  and  leg  color  frequently  differ 
between  sexes;  hence  the  two  sexes  of  some 
species  have  been  described  under  different 
names. 

I  have  freely  used  raw  measurements  in  my 
descriptions,  recognizing  nonetheless  that  they 
are  of  limited  use  in  this  group.  Proportional 
measurements  are  generally  more  useful,  and  I 
include  them  in  my  descriptions  and  diagnoses. 
The  standards  for  these  proportional  measure- 
ments are  head  height  and  length  of  the  first  an- 
tennal  segment — both  measurements  which  are 
proportional  to  general  size. 

This  study  is  limited  to  dried  adult  flies,  so 
structural  characters  are  the  only  ones  which 
form  the  basis  of  my  classification.  What  follows 


WILDER:  NEARCTIC  PTILODEXIA 


is  a  brief  discussion  of  the  taxonomic  impor- 
tance of  the  various  physical  characters  and  how 
their  states  are  determined.  The  headings  and 
general  organization  are  similar  to  those  used  by 
Crosskey  (1973a).  It  is  hoped  that  other  workers 
in  the  Tachinidae  will  adopt  the  same  format, 
eventually  bringing  some  order  to  the  study  of 
variation  in  the  family. 

Body  Color  and  Vestiture 

The  general  integumental  color  of  Ptilodexia 
adults  is  a  dull  brown,  although  adults  of  certain 
more-advanced  species  may  be  black  or  testa- 
ceous, and  teneral  specimens  are  generally  pal- 
er. The  color  of  the  scutellum  compared  to  that 
of  the  rest  of  the  notum  sometimes  is  specifically 
useful  (e.g.,  P.  planifrons-P.  contristans);  how- 
ever, in  adults  of  some  species  (P.  rufipennis), 
it  also  varies  intraspecifically.  The  color  of  the 
abdomen  varies  from  reddish  or  testaceous  with 
a  dark  longitudinal  stripe,  to  concolorous  black 
to  testaceous.  Although  abdominal  color  may  be 
of  occasional  taxonomic  value,  it  almost  always 
varies  between  males  and  females  of  the  same 
species.  In  the  female  it  is  frequently  concolor- 
ous, with  the  longitudinal  stripe  indistinct  or  ab- 
sent. General  body  color  sometimes  varies  cli- 
nally,  and  in  some  species,  smaller,  darker 
populations  exist  in  the  northern  parts  of  the 
range. 

Vestiture  characters  can  aid  in  distinguishing 
members  of  different  species.  These  characters 
seem  to  vary  independently  of  integumental  col- 
or. Facial  tomentum  varies  from  extremely 
heavy — totally  obscuring  the  underlying  integ- 
umental color-to  fine  and  sparse.  Occasionally 
there  is  a  pattern  or  spot  of  color  in  this  vestiture 
which  can  be  distinct  for  a  species  (e.g.,  P.  con- 
tristans,  P.  canescens).  Facial  tomentum  may 
be  dull  (P.  westi)  or  strongly  shining  (P.  incerta). 
The  color  of  the  facial  tomentum  varies  intra- 
specifically. 

The  tomentum  on  the  pleuron  is  of  little  taxo- 
nomic value,  and  that  on  the  notum  is  only 
slightly  more  useful.  The  notal  tomentum  on 
adults  of  some  species  (e.g.,  P.  westi,  P.  arida) 
is  heavy,  abundant,  and  almost  flocculent,  near- 
ly obscuring  the  integumental  color,  whereas  on 
those  of  others  (P.  mathesoni)  it  is  so  fine  that 
the  notum  appears  polished.  In  members  of  oth- 
er species  (P.  conjunct  a),  the  tomentum  is  flat- 
tened and  shiny,  giving  the  notum,  or  parts  of 
it,  a  coppery  hue.  Usually  notal  tomentum  is 


arranged  in  longitudinal  stripes,  but  this  striping 
varies  among  individuals  and  is  of  little  diagnos- 
tic value.  Notal  tomentum  also  varies  between 
the  sexes,  usually  being  heavier  in  the  female. 
Tomentum  on  the  mediotergite  can  be  fine  or 
heavy;  in  specimens  of  P.  agilis  and  P.  mathe- 
soni, however,  it  is  absent. 

Abdominal  tomentum  varies  more  between 
species  than  between  sexes.  It  is  generally  gray- 
ish and  tessellate,  although  in  some  individuals 
it  may  be  gold  or  brownish;  and  it  may  be  shin- 
ing or  dull.  Only  in  adults  of  one  species  (P. 
mathesoni)  has  the  tessellate  pattern  been  re- 
placed by  a  more  uniform  distribution  of  tomen- 
tum, and  even  then,  only  in  the  males.  In  mem- 
bers of  a  few  species  (e.g.,  P.  pacifica,  P. 
ponderosa),  the  grayish  tomentum  is  the  only 
vestiture  on  the  abdomen,  but  in  most  there  are 
patterns  of  brown  or  gold  tomentum  which  can 
aid  in  distinguishing  species.  In  P .  rufipennis 
adults  the  bases  of  the  median  marginal  setae  on 
the  third  and  fourth  abdominal  tergites  are  sur- 
rounded with  gold  tomentum;  adults  of  P.  agilis 
have  a  marginal  band  of  it  on  the  third  and  fourth 
tergites;  those  of  P.  arida  have  brown  tomentum 
overlying  the  longitudinal  stripe. 

These  characters  of  general  color  and  vesti- 
ture show  a  greater  degree  of  variation  within 
Ptilodexia  than  within  all  other  Nearctic  Pro- 
senini,  with  the  possible  exception  of  the  most 
closely  related  genus,  Mochlosoma. 

When  examining  specimens  for  colors  and 
patterns  of  tomentum,  it  is  imperative  that  they 
be  viewed  from  several  different  angles.  Often 
a  pattern  can  be  distinguished  only  if  it  is  seen 
obliquely. 

Chaetotaxy  and  Hairing 

Chaetotaxy  is  of  minor  taxonomic  use  in 
Ptilodexia,  as  it  varies  widely  within  species  but 
little  between  them.  Frequently,  numbers  and 
even  the  presence  of  setae  vary  from  one  side 
of  the  specimen  to  the  other.  This  phenomenon 
is  represented  in  the  descriptions  by  separating 
the  two  states  with  a  slash  (1/0).  Hairs  differ 
from  setae  in  being  much  smaller  and  finer. 

The  taxonomically  useful  setae  on  the  head 
are  the  oral  vibrissae  (Fig.  1).  There  is  usually 
one  pair  (two  in  specimens  of  P.  contristans). 
The  size  and  spatial  relationships  between  the 
vibrissae  and  the  peristomal  setae  can  aid  in 
identifying  adults  in  some  species.  In  P.  rufi- 
pennis adults  the  peristomals  immediately  below 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


frontal  villa 
frontal  seta 

arista 
vlbrlssa 

eplstome 

perlstomal  seta 
haustellum 


ninth   terglte 


surstylus 


poslorbltal 
seta 


adeagus 


FIGURES  1-3.  Structure  of  generalized  Ptilodexia.  Fig.  1. 
Head,  anterolateral  view.  Fig.  2.  Head,  lateral  view.  Fig.  3. 
Genitalia,  lateral  view. 


the  vibrissae  are  short,  becoming  longer  with 
distance  from  the  vibrissae;  in  P.  conjuncta 
adults,  on  the  other  hand,  the  peristomals  are 
subequal  and  nearly  as  long  as  the  vibrissae. 
Numbers  of  peristomal  setae  vary  between  and 
within  species,  but  considerable  overlap  be- 
tween species  is  common.  The  number  and  size 
of  frontal  setae  vary,  but  these  are  even  less 
reliable  characters  than  are  the  number  and  size 
of  the  peristomal  setae. 

The  ocellar,  postocellar,  internal  vertical,  and 
external  vertical  setae  show  some  intraspecific 
differences  in  size  and  number.  I  have  described 
these  differences,  but  they  are  too  variable  to  be 
used  diagnostically.  The  postorbital  setae  vary 
in  length  between  species,  but  not  as  much  as 
between  the  sexes.  The  hairs  which  are  some- 
times inserted  between  them  have  minor  signif- 
icance. In  adults  of  some  species  the  postorbit- 
als  are  long  and  closely  spaced,  while  in  those 
of  others  they  are  interspersed  with  fine  setae 


half  the  length  of  the  postorbitals,  and  in  still 
others  they  are  interspersed  with  tiny  hairs. 

Another  group  of  taxonomically  useful  hairs 
on  the  head  are  those  immediately  ventral  to  the 
postorbitals.  Members  of  species  such  as  P. 
planifrons  and  P.  mathesoni  are  characterized 
as  having  two  to  four  irregular  rows  of  dark  hairs 
between  the  postorbital  setae  and  the  yellow  or 
white  occipital  hair.  Members  of  other  species 
have  only  one  row  of  these  dark  hairs  (P.  cali- 
fornica),  and  those  of  others  have  none  or  just 
a  few  scattered  hairs  (P.  maculata).  This  char- 
acter also  shows  much  intraspecific  variation 
and  must  be  used  cautiously. 

Perhaps  the  best  diagnostic  character  in  Ptilo- 
dexia is  the  hairing  of  the  parafacials  (herein  de- 
fined as  the  sides  of  the  head  bounded  by  the 
apex  of  the  second  antennal  segment,  the  oral 
vibrissae,  the  frontal  suture,  and  the  anterior  eye 
margin).  The  presence,  size,  distribution,  and 
color  of  these  hairs  are  extremely  variable  but 
species  specific.  There  is  slight  variation  in  the 
characteristics  of  these  hairs  between  males  and 
females  belonging  to  the  same  species;  that  is, 
the  parafacial  hairs  of  the  female  are  slightly 
sparser,  finer,  and  are  not  inserted  as  far  ven- 
trally  on  the  parafacial  as  they  are  on  the  male. 
These  hairs  may  be  absent  (P.  rufipennis)(  long, 
dark,  and  abundant  (P.  planifrons);  sparse  and 
pale  (P.  halone)',  minute  and  occurring  only  on 
the  upper  anterior  parafacial  (P.  incerta};  strong 
and  concentrated  at  the  lower  edge  of  the  eye 
(P.  canescens)\  strongly  inclined  anteriad  (P. 
contristans)',  or  inclined  ventrad  (P.  harpasa). 
Many  other  combinations  exist.  It  appears  that 
this  character  can  be  of  diagnostic  value  even  in 
the  Tropics,  where  there  are  many  undescribed 
species.  I  have  illustrated  the  character  state  for 
parafacial  hairs  in  every  Nearctic  species  herein 
described  or  diagnosed.  The  nature  of  the  para- 
facial hairs  is  also  of  importance  in  Mochloso- 
ma,  where  they  are  always  present,  but  not  in 
the  other  Nearctic  Prosenini. 

Parafacial-hair  characteristics  appear  fre- 
quently in  the  key  to  species.  When  the  hairs 
are  small  and  pale,  specimens  must  be  examined 
carefully  from  several  angles;  often  it  is  the  bas- 
es of  the  hairs  rather  than  the  hairs  themselves 
which  are  visible. 

The  parafrontal  hairs  are  of  much  less  taxo- 
nomic  value.  They  are  usually  present,  dark, 
and  are  either  sparse  or  abundant. 


WILDER:  NEARCTIC  PTILODEXIA 


Thoracic  chaetotaxy  is  of  little  diagnostic  im- 
portance in  Ptilodexia.  The  numbers  of  such  se- 
tae as  sternopleurals,  notopleurals,  and  posta- 
lars  are  generally  constant  within  the  genus. 
Others  such  as  posthumerals,  presuturals,  ac- 
rostichals,  dorsocentrals,  and  scutellars  vary 
somewhat  between  species,  but  they  also  show 
considerable  intraspecific  variation.  The  number 
of  humeral  setae  and  discal  scutellars  are  more 
constant,  but  must  be  used  in  combination  with 
other  characters  to  aid  identification. 

The  length  and  density  of  hairs  covering  the 
Ptilodexia  thorax  vary  between  species.  These, 
however,  are  difficult  characters  to  divide  into 
easily  defined  states.  Propleural  hairs  do  not  oc- 
cur in  Ptilodexia  adults,  but  are  present  in  mem- 
bers of  several  closely  related  genera.  They  have 
diagnostic  value  at  the  generic  level. 

Another  group  of  hairs  on  the  thoraces  of 
these  flies  is  the  infrasquamal  setulae,  small 
hairs  inserted  beneath  the  point  of  attachment 
of  the  squamae  or  calypters.  In  adults  of  some 
related  genera  in  the  Prosenini,  these  are  always 
absent.  In  some  Ptilodexia  adults  their  absence 
may  be  a  reliable  specific  character  state  (e.g., 
P.  canescens,  P.  maculata},  but  in  others,  their 
absence  carries  less  taxonomic  importance.  Six- 
ty percent  of  the  specimens  of  P.  rufipennis  ex- 
amined had  infrasquamal  setulae,  but  they  were 
present  in  only  twenty  percent  of  P.  incerta 
specimens.  This  character  is  of  equal  value  in 
both  sexes.  When  using  this  character,  one  must 
realize  that  the  'absence'  of  infrasquamal  setulae 
indicates  absence  on  both  sides  of  the  body. 

Hairs  and  setae  on  the  legs  have  little  diag- 
nostic value;  often  the  setal  length  reflects  total 
body  size  more  than  any  specific  difference.  The 
exception  to  this  is  the  length  of  the  antero-  and 
posteroventral  setae  on  the  posterior  leg  of  the 
male,  which  show  species-level  variation.  These 
are  difficult  to  measure,  however,  and  have  not 
been  used  in  this  revision.  Other  setae  on  the 
femora  show  some  taxonomic  potential,  espe- 
cially the  presence  or  absence  of  anterior  setae 
on  the  posterior  femora. 

Numbers  of  abdominal  setae  vary  intraspecif- 
ically,  but  they  usually  vary  around  a  certain 
number  which  can  be  defined  for  some  species. 
The  presence  or  absence  of  median  marginal  se- 
tae on  the  first  syntergite  can  be  a  useful  char- 
acter. 

The  number  of  median  discal  and  median  mar- 


ginal setae  on  the  third  and  fourth  tergites  is 
useful  in  separating  members  of  some  closely 
related  species  (e.g.,  P.  californica  and  P.  pa- 
cifica),  while  in  others  it  shows  considerable  in- 
traspecific variation.  The  presence  or  absence 
of  lateral  discal  setae  on  these  tergites  will  sep- 
arate members  of  distantly  related  species. 

Length  and  density  characters  of  abdominal 
hairs  have  about  the  same  taxonomic  value  as 
those  characters  in  thoracic  hair.  That  is,  they 
differ  and  seem  to  be  constant  among  members 
of  a  species,  but  are  difficult  to  separate  into 
character  states. 

Although  hairing  on  the  genitalia  varies  only 
slightly  between  species,  the  presence  of  strong 
setae  on  the  ninth  tergite  (epandrium)  is  an  ex- 
cellent diagnostic  character  in  adults  of  P.  con- 
tristans  and  P.  westi. 

Head 

Head  characters  in  Ptilodexia  are  of  more  use 
taxonomically  than  characters  of  any  other  part 
of  the  fly.  Included  are  those  of  chaetotaxy, 
which  have  been  discussed  in  the  previous  sec- 
tion. Drawings  of  the  head,  with  the  terms  used 
in  this  paper,  are  presented  in  Figures  1-2. 

The  head,  in  members  of  this  genus,  is  wide 
and  boxlike.  The  parafacials  and  genae  are  wide 
and  covered  with  fine,  dull-lustered  tomentum. 
The  genae  are  usually  reddish,  contrasting  with 
the  whitish  parafacials  and  genal  dilations.  The 
velvety-appearing  frontal  vitta  extends  from  the 
vertex  to  the  frontal  suture.  The  third  antennal 
segment  is  rarely  longer  than  twice  the  length  of 
the  second  and  bears  an  arista  covered  with  long 
fine  hairs.  Between  the  antennae  is  a  raised  ridge 
or  carina  which  does  not  protrude  beyond  the 
antennae.  The  epistome  may  or  may  not  pro- 
trude. Mouthparts  are  similar  to  those  of  other 
calyptrate  flies  with  the  mentum  length  from  0.3 
to  0.8  times  the  head  height. 

The  width  of  the  parafacial  is  of  considerable 
diagnostic  use  in  Ptilodexia.  Ptilodexia  rufipen- 
nis adults  have  narrow  parafacials,  while  in  P. 
conjunct  a  adults  they  are  quite  wide.  This  char- 
acter is  easier  to  evaluate  in  males  than  in  fe- 
males, and  it  varies  more  among  Ptilodexia 
adults  than  among  Mochlosoma  adults. 

The  shape  of  the  frontal  vitta  can  be  of  use  in 
this  genus.  Below  the  ocellar  triangle  and  be- 
tween the  eyes,  the  frontal  vitta  is  usually  oblit- 
erated (Fig.  4b),  the  parafrontals  becoming  con- 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


FIGURES  4-6.  Comparison  of  specimens  of  Ptilodexia. 
Fig.  4.  Comparison  of  adults  of  two  generalized  species  of 
Ptilodexia  showing  variation  in  width  of  epistome,  position  of 
oral  vibrissae,  and  width  of  frontal  vitta;  A.  head,  anterior 
view,  with  frontal  vitta  not  obliterated  and  distance  of  oral 
vibrissae  from  oral  margin  greater  than  distance  between  oral 
vibrissae;  B.  head,  anterior  view,  with  frontal  vitta  obliterated 
and  distance  of  oral  vibrissae  from  oral  margin  less  than  dis- 
tance between  oral  vibrissae.  Fig.  5.  Comparison  of  antennae 
of  adults  of  two  generalized  species  of  Ptilodexia;  A.  antenna, 
showing  length  of  plumosity  on  arista  shorter  than  length  of 
second  antennal  segment;  B.  antenna  showing  length  of  plu- 
mosity on  arista  longer  than  length  of  second  antennal  seg- 
ment. Fig.  6.  Ptilodexia  ponderosa  (Curran),  holotype,  head 
of  female,  lateral  view. 


tiguous.  In  members  of  some  species  (e.g.,  P. 
canescens,  P.  halone),  however,  the  parafron- 
tals  do  not  touch,  and  the  frontal  vitta  is  contin- 
uous from  the  antennal  base  to  the  ocellar  tri- 
angle (Fig.  4a).  This  character  varies  to  a  similar 
degree  in  specimens  of  Mochlosoma. 

The  size  and  shape  of  the  carina  are  useful  at 
the  supraspecific  level.  Among  Ptilodexia 
species,  the  carina  is  sometimes  distinctly 
shaped  (e.g.,  P.  planifrons).  Carina  characters 
vary  more  among  Mochlosoma  than  Ptilodexia 
adults,  in  general  being  wider  and  better  devel- 
oped in  individuals  of  the  former  genus.  In  mem- 
bers of  several  closely  related  genera  such  as 
Diner  a  and  Hesperodinera,  the  carina  is  strong- 


ly developed,  protruding  from  between  the  an- 
tennae, appearing  almost  bulbous,  and  visible 
from  the  lateral  aspect.  In  Rhamphinina  adults 
it  is  short,  narrow,  and  strongly  keeled.  The  ca- 
rina shows  no  sexual  dimorphism  in  size  and 
shape,  and  except  in  a  few  cases  it  is  species 
specific. 

The  oral  vibrissae  and  adjoining  areas  hold 
characters  of  taxonomic  value  in  Ptilodexia. 
These  are  the  width  of  the  depression  between 
the  bases  of  the  oral  vibrissae  and  the  distance 
of  the  vibrissae  from  the  epistome  (sclerotized 
oral  margin).  Members  of  some  species  of  Ptilo- 
dexia have  this  area  wide  and  short;  an  example 
is  P.  rufipennis,  in  adults  of  which  the  vibrissae 
are  far  apart  and  close  to  the  oral  margin  (Fig. 
4b).  The  opposite  state  is  shown  in  P.  prexaspes 
adults,  which  are  characterized  by  vibrissae  that 
are  far  from  the  oral  margin,  but  not  far  apart 
(Fig.  4a).  In  Rhamphinina  specimens,  the  area 
is  four  times  as  high  as  wide.  This  character  is 
diagnostic  in  a  few  species,  but  in  others  it  varies 
intraspecifically.  It  shows  no  sexual  dimor- 
phism. In  evaluating  this  character,  physical 
measurement  is  necessary;  estimate  is  inade- 
quate. The  invisible  line  connecting  the  vibrissae 
should  pass  through  the  center  of  their  bases, 
and  the  sclerotized  margin  of  the  epistome 
should  serve  as  the  ventral  boundary.  Width  is 
measured  only  at  the  vibrissae;  height  is  mea- 
sured mesially. 

The  epistome  generally  protrudes  in  Ptilo- 
dexia and  Mochlosoma  specimens,  and  al- 
though both  genera  show  variation,  in  Ptilodexia 
adults  it  is  sometimes  species  specific.  In  adults 
of  species  like  P.  prexaspes,  P.  canescens,  and 
P.  halone,  the  oral  margin  projects  slightly  if  at 
all.  As  a  result,  the  lower  anterior  portion  of  the 
head  is  vertical  in  profile  (Fig.  71),  and  in  some 
cases  the  anterior  margin  of  the  head  protrudes 
further  anteriad  at  the  antennae  than  at  the  vi- 
brissae. Ptilodexia  conjunct  a  adults  show  the 
opposite  state,  the  epistome  projecting  strongly, 
as  does  the  lower  anterior  portion  of  the  face 
(Fig.  26).  In  other  prosenines  the  character 
shows  less  intraspecific  variation  than  it  does 
among  species  of  Ptilodexia. 

The  length  of  the  haustellum  is  an  extremely 
valuable  taxonomic  character  in  Ptilodexia  (and 
Mochlosoma},  and  it  is  also  the  chief  difference 
distinguishing  Ptilodexia  from  Mochlosoma 
specimens.  In  individuals  of  the  former  genus, 
the  length  of  the  haustellum  varies  from  0.3  to 


WILDER:  NEARCTIC  PTILODEXIA 


0.9  times  the  head  height,  and  the  shape  is  broad 
and  linear  or  slightly  tapered;  it  is  rigid  in  all 
individuals.  Mochlosoma  specimens  have  the 
haustellum  much  longer  than  the  head  height, 
and  narrow  and  flexible.  In  other  Prosenini  this 
character  serves  to  separate  genera.  In  Ptilo- 
dexia  I  have  compared  the  length  of  the  haus- 
tellum with  the  head  height  and  used  the  result- 
ing ratio  as  a  diagnostic  character  which  varies 
consistently  between  species,  little  within 
species,  and  not  at  all  between  the  sexes.  When 
using  this  character  in  the  key,  actual  measure- 
ments must  be  made;  estimating  the  ratio  is  dif- 
ficult because  a  slender  haustellum  appears  lon- 
ger than  a  broad  one  of  the  same  length. 

There  is  intraspecific  variation  in  the  length  of 
the  haustellum  among  members  of  a  few  species. 
Among  P.  rufipennis  (as  well  as  P.  arida,  P. 
carolinensis,  and  P.  pacified)  specimens,  the 
haustellum  length  varies  locally.  The  length  can 
be  short  in  members  of  one  population  and  no- 
ticeably longer  in  those  of  another.  The  char- 
acter is  still  useful,  though,  since  the  variation 
remains  within  easily  expressed  values. 

The  length  and  shape  of  the  palpi  vary  slightly 
between  members  of  different  species  of  Ptilo- 
dexia.  The  length  is  expressed,  in  this  paper,  as 
a  fraction  of  the  haustellum  length.  Some  Pro- 
senini, such  as  Prosena  and  Senostoma  speci- 
mens, have  short  stubby  palpi;  and  in  Atelog- 
lossa  adults  they  are  completely  absent.  This 
character  varies  among  Mochlosoma  specimens 
much  as  it  does  among  those  of  Ptilodexla.  In 
members  of  P.  arida  and  P.  prexaspes,  the 
length  of  the  palpi  may  nearly  equal  the  length 
of  the  haustellum,  while  in  those  of  P.  obscura, 
it  is  rarely  more  than  0.3  times  the  haustellum 
length. 

The  antennae  possess  some  useful  taxonomic 
characters:  length,  shape,  and  arista  plumosity. 
The  length  of  the  third  segment  is  herein  ex- 
pressed in  terms  of  its  relationship  to  the  rela- 
tively constant  second  segment.  Measurement 
of  the  second  segment  is  taken  from  a  slightly 
anterodorsal  aspect  and  is  the  longest  dorsoven- 
tral  length  of  the  segment. 

Among  Prosena  and  Senostoma  adults, 
length  of  the  third  segment  is  approximately 
twice  the  length  of  the  second;  in  those  of  most 
other  Prosenini,  it  is  considerably  shorter. 
Among  Mochlosoma  species  the  length  varies, 
the  most  usual  state  being  the  third  segment 
equal  to  1.4  to  1.5  the  second.  The  same  is  true 


in  Ptilodexia  species,  where  this  character  can 
be  used  to  separate  adults  of  some  species. 
Ptilodexia  sabroskyi  adults  have  a  short  third 
segment,  subequal  to  or  shorter  than  the  second, 
whereas  those  of  P.  rufipennis  have  the  third 
segment  up  to  twice  the  length  of  the  second. 
Specimens  of  P.  obscura  sometimes  have  the 
third  antennal  article  broadened  apically  instead 
of  slightly  pointed  as  it  is  in  members  of  most 
species. 

The  length  of  the  plumosity  on  the  arista  is  an 
excellent  diagnostic  character  in  Ptilodexia.  I 
have  expressed  it  in  relation  to  the  length  of  the 
second  antennal  segment.  The  arista,  including 
the  plumosity,  is  measured  at  its  greatest  width 
(Fig.  5).  In  specimens  of  P.  rufipennis  and  P. 
harpasa,  two  species  with  long  third  antennal 
segments,  the  length  of  the  plumosity  is  greater 
than  twice  the  length  of  the  second  antennal  seg- 
ment, while  in  those  of  P.  planifrons  and  P. 
prexaspes,  the  length  of  the  plumosity  is  less 
than  or  equal  to  the  length  of  the  second  seg- 
ment. This  character  is  especially  useful  in  sep- 
arating adults  of  the  closely  related  P.  califor- 
nica  and  P.  sabroskyi. 

Thorax 

Most  of  the  thoracic  characters  used  in  this 
revision  have  been  discussed  in  the  sections  on 
vestiture  and  chaetotaxy,  the  remaining  ones  are 
those  of  the  mediotergite,  legs,  and  wings. 

Adults  of  Ptilodexia  have  a  typical  calyptrate 
thorax  with  the  mesothorax  highly  developed, 
and  the  prothorax  and  metathorax  reduced.  The 
scutellum  is  small;  ventral  to  it  is  the  bulging 
postscutellum,  which  distinguishes  members  of 
the  family  Tachinidae.  The  pleuron  is  typical  of 
other  calyptrate  flies.  The  propleuron  is  bare, 
although  the  rest  of  the  pleuron  is  beset  with 
fine  hairs  and  numerous  groups  of  setae.  The 
legs  are  long  with  extremely  long  tarsi.  The 
wings  are  also  long,  the  venation  typical  of  ca- 
lyptrate flies. 

The  mediotergite  is  the  oval  arched  area  ven- 
tral to  the  postscutellum.  In  members  of  some 
species  the  mediotergite  is  dorsally  polished, 
although  other  parts  of  it  may  be  tomentose. 
Care  must  be  taken  when  observing  this  char- 
acter, since  on  adults  of  some  species  the  me- 
diotergite has  a  thin  layer  of  tomentum  and  still 
appears  shiny. 

Leg  color  is  a  taxonomically  useful  character. 
In  species  where  the  color  is  similar  in  members 


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of  both  sexes,  pale-colored  legs  are  diagnostic. 
The  color  of  the  tarsi  can  also  separate  members 
of  different  species  (e.g.,  P.  halone  and  P.  prex- 
aspes}.  Ptilodexia  maculata  specimens  are  dis- 
tinguished by  distinct  femoral  patches  which, 
although  present  on  members  of  other  species, 
are  strikingly  evident  on  those  of  P.  maculata. 
Other  species  exhibit  dimorphism  in  leg  color, 
the  males  with  dark  legs,  the  females  with  pale 
legs  (e.g.,  P.  agilis,  P.  arida).  Two  other 
species,  P.  rufipennis  and  P.  pacifica,  have  fe- 
males with  pale  legs  and  males  with  legs  of  var- 
ied color. 

Wing  venation  is  useful  in  distinguishing 
members  of  some  genera  in  this  tribe  (Nimio- 
glossa).  Within  Ptilodexia  (and  Mochlosoma), 
however,  it  is  of  dubious  value.  Wing  color  is 
constant  within  species  and  can  be  used  diag- 
nostically.  Adults  of  P.  contristans  have  the 
wings  distinctly  darkened  basally,  while  those 
of  P.  mathesoni  have  the  entire  wing  darkened. 
The  colors  of  the  squamae,  epaulet,  and  basi- 
costa  also  show  slight  differences  between  mem- 
bers of  certain  species,  but  they  can  be  varied 
among  those  of  others. 

Abdomen  and  Genitalia 

In  Ptilodexia  adults  as  in  most  Nearctic  Pro- 
senini,  the  abdominal  tergites  meet  ventrally, 
entirely  obscuring  the  sternites.  The  first  tergite 
is  actually  composed  of  two  fused  segments;  the 
next  three  tergites — third,  fourth,  and  fifth — are 
conspicuous.  The  sixth  tergite  is  fairly  broad, 
the  edges  not  meeting  ventrally  (but  embracing 
the  fifth  sternite);  it  and  those  remaining  are 
withdrawn  into  the  fifth  tergite.  The  next  two 
tergites  are  fused  and  become  the  seventh  syn- 
tergite,  which  is  fairly  narrow  in  Ptilodexia 
members  (not  much  wider  than  the  epandrium), 
with  its  surface  oriented  posterodorsally,  as  is 
the  epandrium  (the  ninth  tergite).  This  pattern 
is  similar  in  Mochlosoma  members,  but  in  other 
Prosenini  it  is  different.  In  Prosena  adults,  for 
example,  the  ninth  tergite  appears  to  be  fused 
with  the  seventh  and  eighth,  and  in  members  of 
Hesperodinera,  the  fused  seventh  and  eighth 
tergites  are  exposed  and  greatly  enlarged,  the 
surface  facing  posterad,  the  epandrium  forced 
beneath  the  abdomen. 

Abdominal  color  is  varied  intraspecifically  but 
is  still  useful  as  a  key  characteristic  distinguish- 
ing members  of  some  species.  In  the  key  pre- 
sented herein,  when  the  abdomen  is  described 


as  reddish  laterally,  at  least  the  second  and  third 
tergites  (of  the  male — the  character  is  not  as 
consistently  applicable  to  the  female)  have  the 
integument  reddish  or  rufotestaceous  laterally. 
On  specimens  in  which  the  abdomen  is  concol- 
orous  dark  brown  or  gray,  there  may  be  a  slight 
rufescent  cast  along  the  margins  of  the  tergites. 
This  state  should  not  be  confused  with  the  pre- 
vious one,  in  which  the  reddish  color  extends 
from  the  anterior  to  posterior  margins  of  the  ter- 
gites. 

In  specimens  of  Ptilodexia,  the  external  male 
genitalia  (Fig.  3)  have  taxonomic  value.  The 
characters  which  vary  slightly  between  mem- 
bers of  different  species  are  the  shape  of  the 
ninth  tergite,  the  shape  of  the  surstyli,  and  the 
shape  of  the  cerci.  These  characters  are  useful 
in  distinguishing  members  of  a  few  species,  but 
sometimes  vary  more  intra-  than  interspecifical- 
ly.  Only  in  species  with  extraordinarily  modified 
members  (e.g.,  P.  westi,  P.  rufipennis)  can  the 
external  genitalia  be  called  diagnostic,  and  even 
then,  they  must  be  dissected  for  characters  to 
be  examined  properly.  Often  the  genitalia  of 
adults  of  Mochlosoma  and  Ptilodexia  are  iden- 
tical. The  above-mentioned  characters  vary 
greatly  between  specimens  belonging  to  differ- 
ent genera  and  are  of  excellent  supraspecific 
group  characters. 

Internal  genitalia  are  generally  not  useful  in 
separating  members  of  species  of  Ptilodexia. 
The  aedeagus  is  nearly  identical  in  members  of 
this  genus  and  those  of  Mochlosoma .  The  ejac- 
ulatory  apodeme,  however,  is  useful  in  distin- 
guishing specimens  of  some  species  or  species 
groups.  Its  shape  can  be  distinct,  as  in  P.  con- 
tristans, P.  planifrons,  and  P.  rufipennis  mem- 
bers; between  many  of  the  species,  though,  it 
does  not  vary.  Female  genitalia  show  no  striking 
diagnostic  differences,  with  the  notable  excep- 
tion of  the  surface  sculpturing  of  the  spermathe- 
cae  which,  with  high-magnification  studies,  may 
reveal  specific  differences.  The  reproductive 
systems  of  both  sexes  of  Ptilodexia  have  been 
described  by  Townsend  (1938). 

The  larvae  of  Ptilodexia  have  never  been  de- 
scribed, even  though  there  is  a  figure  of  a  mature 
larva  and  the  puparium  in  Davis  (1919).  The  na- 
ture of  the  cephalopharyngeal  skeletons  of  first 
instar  larvae  (from  the  abdomen  of  gravid  fe- 
males) has  been  used  to  separate  species  in  some 
genera  of  tachinids  (Archytas),  but  the  character 
is  of  no  use  in  Ptilodexia.  Greene  (1922)  de- 


WILDER:  NEARCTIC  PTILODEXIA 


11 


scribed  the  puparium  of  an  unknown  species  of 
Ptilodexia  (erroneously  determined  as  P.  tibi- 
alis). 

Of  the  useful  diagnostic  characters,  none 
works  to  separate  members  of  all  species  from 
those  of  all  others.  Most  of  these  characters  are 
of  high  value  in  distinguishing  members  of  the 
derived  species,  but  when  members  of  certain 
primitive  species  (P.  carolinensis,  P.  major)  are 
examined,  they  lose  much  of  their  value  and 
more  characters  must  be  considered  in  making 
identifications. 


PHYLOGENY 

Present  attempts  to  reconstruct  the  phylogeny 
of  a  genus  or  tribe  in  the  Tachinidae  are  based 
on  incomplete  data  and  should  be  considered 
extremely  tentative  at  best.  Characters  used  at 
generic  and  tribal  levels  are  so  unstable  that  con- 
vergence, loss,  and  acquisition  occur  repeated- 
ly. Most  of  the  species,  and  probably  many  of 
the  genera,  are  unknown  or  poorly  defined  on 
a  worldwide  basis.  Host  relationships  are  largely 
unknown. 

For  the  phylogeny  of  Ptilodexia,  Neotropical 
species  and  representatives  of  closely  related 
genera  were  carefully  examined  to  infer  apo- 
morphic  and  plesiomorphic  states.  Character 
matrices  were  then  constructed  and  phylogenet- 
ic  trees  inferred.  This  method  works  well  when 
trying  to  construct  probable  relationships  in 
higher  categories,  but  for  relationships  among 
species  it  is  not  adequate.  This  is  becuase  the 
characters  distinguishing  species  are  generally 
more  unstable  than  those  distinguishing  families 
or  tribes.  Many  specific  characters  can  be  lost 
or  regained  easily. 

Relationships  within  the  Prosenini  can  be  in- 
ferred only  after  examining  members  of  the  tribe 
on  a  world  basis.  I  have  not  had  the  opportunity 
to  do  this.  I  have  seen  a  few  representatives  of 
the  North  American  genera,  none  of  the  exclu- 
sively Neotropical  genera,  one  of  an  Australian 
genus,  and  one  of  Prosena,  a  worldwide  genus. 
Most  of  the  species  in  these  genera  can  not  yet 
be  identified  with  existing  keys. 

The  characters  used  to  infer  relationships  be- 
tween genera  are  facial  carina,  space  between 
vibrissae,  propleural  hairs,  and  haustellum 
length.  The  form  of  the  facial  carina  provides  a 
good  generic  character,  much  as  it  does  in  the 
tribe  Rutiliini  (Crosskey  1973a).  In  members  of 


Prosenini,  the  carina  was  lost  once,  although 
slight  expression  is  common  in  members  of 
some  species  of  Ptilodexia  and  Mochlosoma. 

The  area  between  the  vibrissae  is  another  sta- 
ble generic  character.  In  Ptilodexia  and  Moch- 
losoma adults,  and  in  those  of  some  of  the  other 
genera,  this  area  is  depressed  and  may  be  flat  or 
slightly  concave,  while  in  members  of  the  more 
primitive  genera,  it  is  slightly  to  strongly  con- 
vex. In  the  primitive  genera,  the  oral  vibrissae 
are  situated  at  or  slightly  above  the  oral  margin, 
while  in  Ptilodexia,  Mochlosoma,  and  Rham- 
phinina  adults,  they  are  inserted  distinctly  dor- 
sal to  the  epistome.  In  Arctophyto  and  Milada 
adults,  their  placement  is  intermediate  between 
that  in  members  of  the  primitive  and  the  derived 
genera. 

Propleural  hairs  are  lacking  in  members  of 
Sentstoma  and  Prosena,  two  of  the  more  prim- 
itive genera.  They  have  been  lost  in  Hespero- 
dinera  adults  and  are  never  seen  in  those  of  the 
advanced  genera. 

The  haustellum  is  longer  than  the  head  height 
in  specimens  of  Prosena,  Prosenoides,  Nimio- 
glossa,  and  Mochlosoma,  but  is  much  shorter 
in  those  of  the  other  genera.  Also,  there  is  great 
variation  in  haustellum  length  among  Ptilodexia 
species  (from  0.3  to  0.9  times  head  height). 

The  position  of  Ptilodexia  within  the  Prosen- 
ini is  advanced.  Assuming  an  Oriental  center  of 
origin  for  the  group,  there  is  a  wealth  of  forms 
(members  of  which  have  a  broad,  blunt,  facial 
carina)  in  the  Oriental  and  Palearctic  regions. 
Some  of  these,  such  as  Prosena  and  Dinera,  are 
widespread.  It  was  probably  a  form  similar  to 
Dinera,  Milada,  and  Arctophyto  which,  along 
with  its  hosts,  crossed  the  Bering  Land  Bridge 
during  favorable  conditions  in  the  late  Tertiary. 
Subsequent  radiation  before  and  during  the 
Pleistocene  must  have  been  great,  for  Ptilodexia 
is  the  most  diverse  genus  of  prosenines  in  the 
western  hemisphere. 

Dinera  and  Arctophyto-Milada  remain  on 
both  sides  of  the  Pacific  with  relatively  few 
species.  Other  small  genera  which  proably  orig- 
inated from  this  complex  are  Ateloglossa,  Dol- 
ichocodia,  Prosenoides,  and  Hesperodinera, 
members  of  which  have  retained  the  inflated  ca- 
rina, and  Myoceropsis,  Rhamphinina,  and  Nim- 
ioglossa,  whose  members  have  lost  it. 

Ptilodexia  and  Mochlosoma,  both  large  gen- 
era, probably  had  similar  origins,  Mochlosoma 
from  a  more  restricted  ancestral  line.  Although 


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the  strong  carina  has  been  lost,  it  is  expressed 
to  a  moderate  degree  in  members  of  some 
species  in  both  genera.  It  is  my  opinion  that  the 
only  character  which  separates  Ptilodexia  from 
Mochlosoma  adults,  the  haustellum  shape  and 
length,  is  a  phylogenetically  sound  one,  that  is, 
its  origin  in  Mochlosoma  is  monophyletic.  Al- 
though radiation  of  both  genera  has  been  great, 
divergence  between  Mochlosoma  and  Ptilo- 
dexia members  is  only  slight. 

Ateloglossa  and  Hesperodinera  had  their 
origins  from  a  Dinera-like  ancestor,  members  of 
the  former  having  lost  the  palpi  and  the  latter 
the  propleural  hairs,  but  both  having  retained 
the  squarish  head  and  inflated  facial  carina.  Pro- 
senoides  adults,  on  the  other  hand,  bear  a  closer 
resemblance  to  Prosena  specimens,  and  the  two 
may  be  closely  related.  Both  Nimioglossa  and 
Rhamphinina  had  their  origins  early  in  the  Ptilo- 
dexia-Mochlosoma  line,  their  members  having 
diverged  from  the  ancestral  forms  in  having  the 
area  between  the  oral  vibrissae  depressed  and 
the  vibrissae  placed  considerably  above  the  oral 
margin,  as  do  representatives  of  Ptilodexia  and 
Mochlosoma. 

Within  Ptilodexia,  characters  indicating  rela- 
tionship are  difficult  to  determine.  The  ancestral 
and  derived  states  of  a  few  of  these  characters 
have  been  inferred. 

Small  size  and  dark  color  seem  to  be  primitive 
states  within  Ptilodexia,  while  the  derived  states 
are  large  size  and  pale  color.  The  primitive  state 
of  the  length  of  the  third  antennal  segment,  the 
length  of  the  plumosity  on  the  arista,  and  length 
of  the  haustellum  is  an  intermediate  one,  with 
the  derived  states  being  short  and  long.  The 
presence  of  both  parafacial  hairs  and  infrasqua- 
mal  setulae  appear  to  be  primitive  within  this 
genus.  Primitively  the  oral  margin  is  narrow  and 
projecting,  while  the  more  advanced  forms  show 
it  to  be  wide  and  not  projecting.  The  primitive, 
typical  shape  of  the  ejaculatory  apodeme  can  be 
seen  in  most  species  (Fig.  44).  Members  of  some 
of  the  advanced  species  have  this  structure  mod- 
ified in  various  ways  (Fig.  29,  35),  although 
those  of  others  do  not. 

The  genus  Ptilodexia  in  North  America  is 
comprised  of  six  loosely  knit  groups.  The  first, 
and  probably  most  primitive  of  these,  is  the 
agilis  group,  which  consists  of  P.  agilis,  P.  ob- 
scura,  and  P.  mathesoni.  The  next  group  is  the 
carolinensis  group,  with  member  species  P. 
carolinensis,  P.  halone,  P.  prexaspes,  and  P. 


canescens.  The  harpasa  group  is  composed  of 
P.  rufipennis,  P.  arida,  P.  harpasa,  and  P.  pon- 
der osa.  In  the  major  group  are  P.  major,  P. 
incerta,  P.  maculata,  and  P.  flavotessellata. 
The  conjuncta  group  contains  P.  conjuncta,  P. 
planifrons,  P.  contristans,  and  P.  westi;  and  the 
californica  group,  P.  californica,  P.  sabroskyi, 
and  P.  pacifica. 

The  character  states  which  segregate  mem- 
bers of  these  groups  are  vague  and  difficult  to 
define,  but  since  the  groups  appear  to  have  both 
a  zoogeographical  and  morphological  basis,  they 
will  be  discussed.  Their  relationships  to  each 
other  are  somewhat  less  clear. 

Members  of  the  agilis  group  are  small,  dark 
flies  with  little  red  color  on  the  abdomen  and  a 
short  haustellum.  The  legs  of  the  females  of  P. 
agilis  and  P.  mathesoni  are  pale,  while  those  of 
P.  obscura  are  dark.  The  mediotergite  is  pol- 
ished or  shiny  in  members  of  all  three  species. 
This  is  probably  the  most  primitive  group  of 
Ptilodexia;  P.  agilis  members  perhaps  being 
similar  to  those  of  the  prototype  of  the  genus. 
Ptilodexia  agilis  is  a  widespread  western  form, 
occurring  into  central  Texas;  P.  mathesoni, 
closely  related,  is  a  northern  form  found  in  New 
York,  Michigan,  and  eastern  Canada.  Ptilodexia 
obscura  has  a  distribution  from  the  Appalachi- 
ans to  the  Rocky  Mountains  and  is  nearly  com- 
plimentary to  that  of  P.  agilis. 

Species  included  in  the  carolinensis  group 
have  members  with  a  nonprojecting  epistome 
and  a  nearly  vertical  anterior  head  profile.  All 
adults  have  relatively  short  plumosity  on  the 
arista  and  the  abdomen  reddish  laterally.  Two  of 
these  species,  P.  halone  and  P.  prexaspes,  have 
limited  east  coast  distributions.  Ptilodexia  ca- 
nescens occurs  across  the  northern  United 
States  and  Canada,  while  P.  carolinensis  is 
widely  distributed  from  the  east  coast  to  the 
Rocky  Mountains.  Ptilodexia  carolinensis  is 
probably  the  oldest  of  the  four,  P.  canescens, 
P.  halone,  and  P.  prexaspes  being  derived  from 
it  during  the  Pleistocene. 

The  harpasa  group  is  the  most  ambiguous  of 
all,  containing  most  of  the  Antillean  and  many 
Mexican  species.  Members  of  these  species  all 
have  long  antennae  and  long  plumosity  on  the 
arista;  some  lack  parafacial  hairs.  Ptilodexia  ru- 
fipennis occurs  from  the  east  coast  to  the  Rocky 
Mountains  and  across  Canada;  P.  harposa  is 
more  restricted,  P.  arida  is  restricted  to  the 
Southwest  and  Mexico,  while  P.  ponder  osa  is 


WILDER:  NEARCTIC  PTILODEXIA 


13 


CALIFORNICA         CONJUNCTA       AGILIS 


OBSCURA       CAROLINENSIS        MAJOR       HARPASA 


FIGURE  7.     Inferred  phylogeny  of  Ptilodexia  species  groups. 


probably  a  West  Indian  species,  with  one  record 
from  Florida.  Assuming  that  P.  harpasa  is  the 
closest  to  the  ancestor  of  the  group,  P.  rufipen- 
nis  became  the  most  widespread  and  P.  arida 
and  P.  ponderosa  radiated  in  the  southern  lati- 
tudes. 

Ptilodexia  major,  the  most  primitive  member 
of  the  next  group  shows  slight  similarities  to 
specimens  of  P.  harpasa.  It  ranges  widely 
throughout  the  Midwest,  Southwest,  and  Mex- 
ico. Ptilodexia  incerta  has  an  eastern  distribu- 
tion almost  exactly  complementary  to  that  of  P. 
major,  while  P.  maculata  and  P.flavotessellata 
are  restricted  in  the  Southwest  and  Midwest. 
These  species  all  have  members  with  short, 
pale,  parafacial  hairs. 

The  next  group,  conjuncta,  is  probably  de- 
rived directly  from  the  ancestral  agilis  group 
and  consists  of  only  western  species.  Ptilodexia 
conjuncta,  its  most  primitive  member,  ranges 
throughout  the  Rocky  Mountains  from  Canada 
into  Mexico  and  west  to  California.  Ptilodexia 
planifrons  and  P.  contristans  extend  from  the 
southwestern  United  States  into  Mexico;  and  P. 
westi,  a  close  relative  of  P.  contristans,  is  re- 
stricted to  the  extreme  southern  Midwest  and 
the  Southwest. 


The  last,  or  calif ornica  group,  is  related  to  the 
conjuncta  group  and  probably  had  a  similar  or- 
igin. Its  three  species  are  confined  to  California 
and  the  West  Coast,  and  members  of  these  taxa 
show  similarities  only  to  members  of  the  con- 
juncta group. 

A  graphic  representation  of  the  relationships 
of  these  species  groups  is  given  in  Figure  7.  Pos- 
sible events  at  the  numbered  branching  points 
are  as  follows: 

1.  Major  east-west  split.  Eastern  group  mem- 
bers with  parafacial  hairs  extending  low  on 
the  face;  western  group  members  with 
parafacial  hairs  high  on  the  face. 

2.  Widening  of  face;  lighter  general  color. 
Some  females  with  yellow  legs;  lengthen- 
ing of  plumosity  on  arista.  Some  advanced 
members  radiating  widely;  radiation  into 
Mexico  and  Antilles. 

3.  Lengthening  of  haustellum,  increase  in 
size,  widening  of  face.  Extensive  radiation 
into  Mexico. 

4.  Slight  increase  in  size.  Flattening  of  oral 
margin  and  anterior  facial  margin;  short- 
ening of  haustellum  and   plumosity   on 
arista. 


14 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


5.  Shortening  of  haustellum.   Radiation  and 
isolation  in  southwestern  United  States 
and  Mexican  Pleistocene  refugia. 

6.  Pleistocene  isolation  in  California.   De- 
crease in  abundance  of  parafacial  hairs  and 
length  of  haustellum. 

ZOOGEOGRAPHY 

Although  the  dispersal  powers  of  Diptera  are 
relatively  great,  the  distribution  patterns  seen  in 
Ptilodexia  seem  to  be  dependent  upon  those  of 
their  hosts,  the  Scarabaeidae.  Distribution  of 
some  Scarabaeidae  are  well  known,  and  their 
possible  histories  have  been  discussed  in  several 
papers  (Howden  1963;  1966). 

All  statements  made  in  this  section  are  ten- 
tative. The  patterns  discussed  are  those  of 
species  of  Ptilodexia,  but  interpretations  of 
those  patterns  are  those  which  have  been  of- 
fered for  some  of  the  species  of  Scarabaeidae. 
No  host  specificity  has  been  found,  and  it  is  only 
speculation  that  similar  patterns  of  Ptilodexia 
and  their  scarabaeid  hosts  are  due  to  similar  his- 
tories. 

Howden  (1966)  stated  that  North  American 
species  of  Phyllophoga  show  a  decline  in  num- 
bers from  Georgia  to  Canada  and  from  Texas  or 
Arizona  to  Nebraska.  This  is  true  of  Ptilodexia. 
He  also  stated  that  if  certain  areas  of  Texas  were 
included  with  Arizona  and  New  Mexico,  there 
would  be  little  overlap  between  the  eastern  and 
western  faunas  (approximate  dividing  line,  100th 
meridian).  This  holds  true  for  the  most  part  in 
Ptilodexia.  However,  a  number  of  eastern 
species  occur  all  the  way  into  British  Columbia 
in  the  northern  parts  of  their  ranges.  As  with 
Phyllophaga,  many  of  the  southern  Arizona  rec- 
ords represent  the  northern  limits  of  Mexican 
species. 

When  plotting  the  centers  of  distribution  of 
species  of  Ptilodexia,  it  was  noted  that  six 
species  groups  could  be  defined  geographically. 
These  were  the  same  six  groups  which  had  been 
structurally  and  zoogeographically  defined 
above.  Although  these  groups  show  that  the 
phylogeny  presented  herein  has  a  zoogeograph- 
ical  basis,  they  are  not  the  best  groupings  for 
discussing  zoogeography. 

I  have  categorized  the  species  of  Ptilodexia 
into  six  zoogeographical  groups,  based  on  their 
complete  distributions  rather  than  centers  of  dis- 


tribution. The  relationships  of  these  categories 
may  give  insight  into  the  historical  zoogeogra- 
phy of  the  group. 

The  first  of  these  is  an  extreme  northern  pat- 
tern shown  by  P.  canescens  and  P.  mathesoni. 
The  distribution  is  almost  exclusively  in  areas 
which  were  previously  glaciated.  The  range  of 
P.  mathesoni  (Fig.  17)  is  restricted  to  Michigan, 
New  York,  and  eastern  Canada.  Its  ancestral 
and  most  closely  related  species,  P.  agilis,  oc- 
cupies a  large  area  from  the  Rocky  Mountains 
west,  extending  eastward  into  Texas  (Fig.  12). 
Ptilodexia  canescens,  from  the  Caroline nsis 
group,  inhabits  the  northern  United  States  and 
Canada  from  Newfoundland  to  British  Colum- 
bia. In  the  East  it  extends  southward  only  to  the 
previous  front  of  the  Wisconsin  glaciation,  while 
in  the  West  it  extends  southward  into  eastern 
Idaho,  western  Wyoming,  and  northern  Utah 
(Fig.  67).  The  distribution  of  a  species  in  pre- 
viously glaciated  areas  without  representation 
south  of  the  glacial  front  is  fairly  uncommon 
(Ross  1965).  This  deglaciated  area  may  have  of- 
fered considerable  opportunity  for  expansion  to 
certain  Scarabaeidae  and  their  Ptilodexia  para- 
sitoids. 

The  next  group  has  a  widespread  distribution, 
throughout  the  eastern  United  States  into  the 
plains  states  and,  in  some  cases,  even  further 
west.  Of  these  species,  only  P.  harpasa  (Fig. 
82)  lacks  representation  in  the  lower  Midwest. 
The  other  species,  P.  carolinensis  (Fig.  62),  P. 
incerta  (Fig.  103),  P.  obscura  (Fig.  22),  and  P. 
rufipennis  (Fig.  88)  occur  widely  throughout  the 
Midwest  and  the  East. 

The  remaining  eastern  distribution  pattern  is 
that  of  P.  halone  and  P.  prexaspes.  Both  of 
these  species  belong  to  the  carolinensis  group; 
they  are  closely  related  and  complementary  in 
distribution.  Ptilodexia  prexaspes  occurs  in 
Florida  and  along  the  Atlantic  coast  to  Virginia 
(Fig.  77),  while  P.  halone  is  found  in  Mississip- 
pi, Tennessee,  and  along  the  coast  from  Mary- 
land to  New  York  (Fig.  72).  It  is  possible  that 
these  relatively  uncommon  species  are  host-spe- 
cific parasites  of  some  of  the  large,  flightless 
scarabs  found  in  the  Southeast  and  discussed  by 
Howden  (1963). 

Two  species,  P.  agilis  and  P.  conjuncta,  have 
large  western  ranges,  the  former  from  British 
Columbia  to  Texas  and  west  to  the  Pacific  coast 
(Fig.  12),  the  latter  from  Mexico  to  British  Co- 


WILDER:  NEARCTIC  PTILODEXIA 


15 


lumbia,  west  to  the  coast  (excluding  California), 
and  eastward  through  Canada  to  Ontario  (Fig. 
27). 

Ptilodexia  calif  arnica  (Fig.  47),  P.  pacific  a 
(Fig.  52),  and  P.  sabroskyi  (Fig.  57)  have  ranges 
which  are  restricted  to  the  west  coast  of  the 
United  States.  It  is  possible  that  some  unique 
local  populations  are  parasitic  on  the  large  flight- 
less genera  of  scarabs  (such  as  Pleacama)  which 
survived  in  situ  during  the  Pleistocene. 

The  remaining  distribution  group  is  the  most 
common  in  Ptilodexia,  occurring  in  at  least  sev- 
en species.  This  is  a  southwestern  distribution, 
with  species  which  may  have  had  Mexican  re- 
fugia.  Four  of  these  are  Mexican  species  whose 
ranges  extend  northward  into  the  mountainous 
regions  of  Arizona  and  New  Mexico,  rarely  into 
Utah  and  Idaho.  These  four  are  P.  contristans 
(Fig.  37),  P.  planifrons  (Fig.  32),  P.  maculata 
(Fig.  108),  and  P.  arida  (Fig.  93),  all  recently 
differentiated.  It  is  possible  that  the  ranges  of 
many  other  Nearctic  Mexican  species  also  ex- 
tend into  these  areas,  but  specimens  have  not 
yet  been  taken  by  collectors. 

Two  of  this  southwestern  group,  P.  westi  and 
P.  flavotessellata,  apparently  do  not  range  into 
Mexico.  The  former  occurs  broadly  along  the 
international  boundary  from  central  Arizona  to 
eastern  Texas  and  into  Oklahoma  and  southern 
Kansas  (Fig.  42),  while  P.  flavotessellata  occurs 
in  northern  New  Mexico,  Colorado,  and  Ne- 
braska (Fig.  1 13). 

The  last  species  in  the  southwestern  group  is 
P.  major.  Its  distribution  is  a  combination  of  the 
ranges  of  the  previous  two  groups,  extending 
from  Mexico  (where  it  is  widespread)  into  the 
mountains  of  Arizona,  New  Mexico,  and  Colo- 
rado and  through  Texas  into  the  Plains  in  Ne- 
braska and  Kansas  (Fig.  98). 

Unlike  those  of  Phyllophaga  (Howden  1966), 
eastern  species  of  Ptilodexia  frequently  occur 
from  Georgia  to  southern  Ontario;  others  range 
broadly  across  the  northern  part  of  the  United 
States  and  Canada.  I  see  this  deviation  as  a  re- 
sult of  the  vagility  of  these  flies  and  the  probable 
capability  of  developing  in  different  hosts,  fac- 
tors which  may  account  for  other  deviations 
from  typical  scarabaeid  distributions. 

Inferring  the  historical  zoogeography  of  Ptilo- 
dexia is  extremely  speculative.  The  genus,  as 
we  know  it,  probably  evolved  on  this  continent, 
its  ancestor  reaching  the  area  via  the  Bering 


Land  Bridge  during  the  Tertiary.  By  the  onset 
of  the  Pleistocene,  most  of  the  species  were 
probably  already  established.  Pleistocene  cli- 
matic fluctuations  must  have  affected  the  distri- 
bution patterns  we  see  in  the  genus  today. 

The  eastern  species  in  our  fauna  may  have 
occupied  southeastern  Pleistocene  refugia,  most 
of  them  expanding  westward  in  the  north  after 
the  ice  sheets  retreated.  Ptilodexia  agilis  seems 
to  have  been  much  more  widespread  at  one 
time,  one  of  the  species  derived  from  it  being 
found  only  in  the  Northeast.  While  P.  agilis  may 
have  had  a  wide  refugium,  P.  conjuncta  and  P. 
californica  perhaps  survived  the  Pleistocene  in 
Mexico  and  California,  respectively,  separated 
by  the  extensive  desert  barriers  of  the  time.  The 
three  species  endemic  to  California  were  prob- 
ably separated  from  the  other  species  at  a  rela- 
tively early  time,  closely  resembling  each  other 
considerably  more  than  any  other  species.  Their 
refugia  were  in  central  and  southern  California, 
and  subsequent  recolonization  proceeded  no 
further  north  than  the  southern  limits  of  the  ice 
sheet.  The  southwestern  groups  could  have  sur- 
vived the  glacial  periods  in  situ  or  in  Mexican 
refugia.  It  appears  that  P.  major  was  once  a 
widespread  species,  extending  well  into  Mexico 
before  the  Pleistocene  and  giving  rise  to  many 
species  there. 

The  Southwest,  including  Texas  and  Arizona, 
has  the  largest  number  of  endemics.  In  the 
warmer  parts  of  the  country  such  as  these,  more 
generations  per  year  are  possible,  and  evolution 
can  proceed  at  a  faster  pace  than  in  the  north. 
This  ripay  account,  in  part,  for  the  large  number 
of  endemics;  it  also  helps  explain  the  numerous 
divergent  populations  seen  in  California  and 
Texas  as  well  as  the  tremendous  diversity  of  the 
genus  in  Mexico. 

Genus  PTILODEXIA  Brauer  and  Bergenstamm 

Ptilodexia  BRAUER  AND  BERGENSTAMM,  1889:119  (Type- 
species,  Ptilodexia  carolinensis  Brauer  and  Bergenstamm, 
1889,  by  original  designation.) 

Myoceropsis  TOWNSEND,  1915:23  (Type-species,  Rhynchio- 
dexia  flavotessellata  Walton  by  original  designation.) 

Rhamphinina,  authors,  not  Bigot. 

Rhynchiodexia,  authors,  not  Bigot. 

Rhynchodexia,  emend.  Wulp,  1891. 

The  genus  Ptilodexia  Brauer  and  Bergen- 
stamm is  confined  to  the  New  World.  It  is  best 
represented  in  the  Neotropical  region,  as  is  the 
entire  tribe  Prosenini. 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


DIAGNOSIS. — Members  of  Ptilodexia  can  be 
distinguished  from  those  of  all  closely  related 
species  of  Nearctic  Prosenini  by  the  following 
combination  of  character  states:  propleuron 
bare;  facial  carina  sometimes  well  developed 
(but  never  broad,  blunt,  and  separating  the  an- 
tennae); haustellum  shorter  than  the  head 
height;  apical  cell  open  or  closed  at  wing  margin; 
and  infrasquamal  setulae  usually  present. 

DESCRIPTION. — Color  black  to  reddish,  usu- 
ally with  thorax  dark  and  abdomen  pale  with 
dark  longitudinal  stripe.  Head  with  face  broad 
in  profile,  anterior  margin  usually  vertical;  epi- 
stomal  margin  somewhat  projecting;  frontal  vitta 
strongly  narrowed  between  eyes,  often  obliter- 
ated; facial  tomentum  heavy  to  sparse,  dull  to 
shining,  color  varied,  but  usually  grayish;  para- 
facial  hairs  varied,  absent  or  present;  carina 
long,  not  much  deeper  than  width  of  third  an- 
tennal  segment;  postocular  setae  long;  one  or 
two  pairs  of  oral  vibrissae;  epistome  generally 
projecting  to  some  degree;  haustellum  rigid,  var- 
ied, from  0.3  to  0.8  times  head  height;  palpi  long. 
Antennae  with  length  of  third  segment  varied 
from  one  to  two  times  length  of  second;  arista 
with  length  of  plumosity  more  than  length  of 
second  antennal  segment.  Thorax  with  propleu- 
ron bare;  mesonotum  strongly  or  weakly  tomen- 
tose,  usually  indistinctly  striped;  three  or  four 
pairs  of  presutural  and  postsutural  acrostichals; 
a  tuft  of  small  hairs  on  postalar  wall.  Wing 
length  2.5  times  width;  apical  cell  open  or  closed 
at  wing  margin;  infrasquamal  setulae  present  or 
absent.  Legs  dark,  tibiae  lighter  in  most  cases, 
posterior  tarsi  very  long,  1 .5  times  length  of  tib- 
ia; claws  and  pulvilli  long.  Abdomen  broad, 
conical,  tomentum  usually  in  large  irregular 
patches;  numerous  median  discal  and  marginal 
setae  on  abdominal  tergites;  four  abdominal  ter- 
gites  visible,  lateral  margins  meeting  ventrally. 
Ge nit alia  slightly  withdrawn,  terminal,  axis  ver- 
tical; cerci  and  surstyli  variously  modified.  Fe- 
male differs  from  male  in  following  ways:  frontal 
vitta  wide  with  sides  subparallel;  eyes  widely 
separated;  profrons  a  little  wider;  frontal  and 
peristomal  setae  not  as  abundant;  vertex  with 
few  hairs  or  setae;  postocular  setae  shorter  and 
sparser;  height  of  eyes  distinctly  less;  frontal 
orbital  setae  present  (Fig.  6).  Thorax  with  fewer 
setae  and  hairs  and  more  heavily  tomentose; 
thorax  and  legs  frequently  lighter  in  color  with 
fewer  and  shorter  major  setae  and  hairs;  tarsal 
claws  and  pulvilli  much  shorter.  Abdomen 


broader,  shorter,  and  much  more  heavily  to- 
mentose; usually  with  fewer  median  discals  and 
often  lacking  other  setae;  integumental  color 
uniform  brown  or  gray  in  many  females,  even 
when  it  is  marked  in  males  of  the  same  species. 

Brauer  and  Bergenstamm  erected  the  genus 
Ptilodexia  in  1889  for  the  North  American 
species  P.  carolinensis.  There  has  been  much 
confusion  since  that  time  regarding  the  limits  of 
the  genus.  This  is  because  several  characters 
normally  constant  within  tachinid  genera  vary 
among  Ptilodexia  species.  These  characters  in- 
clude the  presence  or  absence  of  parafacial  hairs 
and  infrasquamal  setulae. 

Prior  to  1889,  Macquart  and  Walker  described 
species  belonging  to  Ptilodexia  in  the  genus 
Dexia  Meigen,  while  Bigot  (1885)  created  the 
genus  Rhamphinina  for  those  species  he  de- 
scribed. Bigot  thought  that  Rhamphinina,  a  neo- 
tropical genus,  and  Rhynchiodexia,  one  of  his 
Australian  genera,  could  be  distinguished  from 
one  another  by  the  presence  or  absence  of  a 
facial  carina.  Wulp  (1891)  considered  this  to  be 
an  inconsistent  character. 

Wulp  (1891)  emended  the  name  to  Rhyncho- 
dexia ,  which  he  used  for  what  we  now  call  Ptilo- 
dexia. He  felt  that  Rhynchodexia  and  Rham- 
phinina were  congeneric  and  mistakenly  placed 
one  Mexican  species,  contristans,  in  Hystri- 
chodexia. 

West  (1924;  1925),  a  North  American  worker, 
thought  that  the  species  with  hairy  parafacials 
belonged  to  Ptilodexia  while  those  with  bare 
parafacials  were  Rhynchodexia.  Austen  (1907) 
shared  this  opinion.  Later,  West  (1950)  agreed 
with  Curran  (1934)  that  the  two  were  probably 
one  and  Rhynchodexia  was  the  proper  name  for 
the  complex.  Reinhard  (1943)  stated  that  the 
name  Ptilodexia  was  available  for  American 
species. 

Examination  of  species  of  Rhynchodexia, 
now  Senostoma  (Crosskey  1973a),  shows  that 
this  genus  differs  from  Ptilodexia  in  having  a 
pronounced  facial  carina  as  well  as  numerous 
other  differences  which  will  be  discussed  later. 
Rhamphinina  dubia,  the  type-species  of  the  ge- 
nus, is  not  a  Ptilodexia. 

The  name  Estheria  tibialis  Robineau-Desvo- 
idy  is  frequently  used  for  species  of  Ptilodexia 
(Townsend  1921;  Aldrich  1905;  Austen  1907). 
The  type  of  this  species  is  lost,  so  we  cannot 
know  if  E.  tibialis  belongs  to  Ptilodexia.  How- 
ever, since  a  characteristic  of  Estheria  is  the 


WILDER:  NEARCTIC  PTILODEXIA 


17 


presence  in  its  members  of  a  petiolate  apical 
cell,  and  since  this  rarely  occurs  in  Ptilodexia, 
I  agree  with  previous  workers  who  have  chosen 
to  reject  the  name  E.  tibialis. 


Key  to  the  Nearctic  Species  of  Ptilodexia 

la.  Parafacial  hairs  present,  although  very 
small  and  pale  in  some  individuals;  legs 
of  females  varied  in  color  2 

Ib.  Parafacial  hairs  absent;  legs  of  females 
pale  in  color  27 

2a.  Infrasquamal  setulae  present 5 

2b.  Infrasquamal  setulae  absent  3 

3a.  Parafacial  hairs  dark,  long,  present  on 
most  of  parafacial  (Fig.  66);  length  of 
plumosity  on  arista  subequal  to  length  of 
second  antennal  segment;  face  with 
traces  or  spots  of  brownish  tomentum 
(northeastern  U.S.,  trans-Canada, 

northern  mountain  states)    

canescens  (Walker) 

3b.  Parafacial  hairs  pale  and/or  short,  pres- 
ent only  on  upper  anterior  portion  of 
parafacial;  length  of  plumosity  on  arista 
at  least  1.5  times  length  of  second  anten- 
nal segment  (Fig.  5b);  facial  tomentum 
concolorous  silvery  gray  or  yellowish  4 

4a.  Femora  of  members  of  both  sexes  brown 
or  black  (eastern  U.S.  to  about  100th 
meridian)  incerta  (West)  (in  part) 

4b.  Femora  of  members  of  both  sexes  or- 
ange with  definite  black  or  brown 
patches  on  flexor  surfaces,  coxae  also 
with  dark  patches  (Arizona  and  New 
Mexico) maculata  n.sp. 

5a(2a).  Flies  pale  colored;  thorax,  abdomen, 
and  femora  pale  brown  to  orange,  or 
width  of  depression  between  oral  vibris- 
sae  less  than  distance  between  oral  vi- 
brissae  and  oral  margin  (Fig.  4a)  6 

5b.  Flies  dark;  thorax,  abdomen,  or  legs 
brown  or  darker  in  color;  width  of 
depression  between  oral  vibrissae  equal 
to  or  greater  than  distance  between  oral 
vibrissae  and  oral  margin  (Fig.  4b)  ..  10 

6a.  Integument  of  tarsi  pale,  concolorous 
with  legs  ventrally  halone  (Walker) 

6b.  Integument  of  tarsi  brown  or  black 7 

7a.  Width  of  the  depression  between  oral 
vibrissae  greater  than  distance  from  vi- 
brissae to  oral  margin  8 

7b.  Width  of  depression  between  oral  vibris- 


sae less  than  or  equal  to  distance  from 

vibrissae  to  oral  margin  (Fig.  4a) 

prexaspes  (Walker) 

8a.  Parafacial  hairs  minute,  confined  to  up- 
per anterior  parafacial;  haustellum  length 
0.6  times  head  height  (Fig.  6)  (southern 

Florida)    ponderosa  (Curran) 

8b.  Parafacial  hairs  long,  scattered  on  para- 
facial; haustellum  length  no  more  than 
0.5  times  head  height  (southwestern 

U.S.  and  Texas) 9 

9a.  Parafacial  hairs  dark,  coarse  and  abun- 
dant (Fig.  61);  femora  or  notum  brown 
in  many  individuals;  this  color  form  un- 
common (Texas)  

____  carolinensis  Bauer  and  Bergenstamm 

(in  part) 

9b.  Parafacial  hairs  pale  or  light  brown,  fine, 
sparse  (Fig.  112);  femora  and  notum  pale 
orange-brown  (southwestern  U.S.  into 

Colorado  and  Nebraska)    

flavotessellata  (Walton) 

10a(5b).  Mediotergite  polished  immediately 
beneath  postscutellum;  parafacial  hairs 
distant  from  eye;  haustellum  0.5  times 
head  height  or  less  (Fig.  11);  legs  of  fe- 
males yellow;  species  with  members 
small,  dark  11 

lOb.  Mediotergite  with  at  least  a  fine  dusting 
of  tomentum;  parafacial  hairs  and  haus- 
tellum varied  12 

lla.  Abdomen  and  thorax  black,  strongly 
shining;  abdomen  with  tomentum  evenly 
distributed;  parafacial  hairs  black,  coarse 
(Fig.  16);  squamae  of  males  dark  brown 

(northcentral  and  eastern  U.S.) 

mathesoni  (Curran) 

lib.  Abdomen  with  tomentum  in  large  irreg- 
ular patches;  thorax  with  definite  tomen- 
tose  striping;  parafacial  hairs  black,  fine 
(Fig.  1 1);  squamae  of  males  white  to  pale 

brown  (western  U.S.  and  Texas) 

agilis  Reinhard  (in  part) 

12a(10b).  Length  of  plumosity  on  arista  less 
than  or  equal  to  1.25  times  length  of  sec- 
ond antennal  segment  (southwestern 
U.S.  and  Mexico) 13 

12b.  Length  of  plumosity  on  arista  more  than 
1.25  times  length  of  second  antennal  seg- 
ment (widespread) 15 

13a.  Abdomen  and  scutellum  concolorous 
dark  gray  or  brown,  slightly  lighter  on 
ventral  margins  of  tergites  in  a  few  in- 


18 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


dividuals:  ninth  tergite  and  parafacial 
hairs  varied;  haustellum  length  0.7  to  0.8 
times  head  height  (Fig.  31)  14 

13b.  Abdomen  and  scutellum  distinctly  red- 
dish laterally;  ninth  tergite  with  several 
strong  setae  (Fig.  33);  parafacial  hairs 
long,  dark,  occurring  along  parafacial 
ventrally  to  level  of  oral  vibrissae;  haus- 
tellum length  0.6  times  head  height  (Fig. 
36)  contristans  (Wulp) 

14a.  Parafacial  hairs  long,  coarse,  numerous, 
occurring  on  entire  parafacial  (Fig.  31) 
(Arizona,  New  Mexico  into  Mexico)  .. 
planifrons  (Wulp) 

14b.  Parafacial  hairs  fine,  sparse,  occurring 
only  on  anterior  parafacial  (Fig.  41) 

(Texas,  Oklahoma,  Kansas)  

westi  n.sp.  (in  part) 

15a(12b).  Parafacial  hairs  short,  pale  or  oth- 
erwise inconspicuous  16 

15b.  Parafacial  hairs  long,  dark,  although 
sparsely  placed  in  some  individuals  __  20 

16a.  Mediotergite  polished;  legs  of  females 
yellowish;  haustellum  length  no  more 
than  0.5  times  head  height  (Fig.  1 1);  scu- 
tellum concolorous  with  rest  of  notum 
agilis  Reinhard  (in  part) 

16b.  Mediotergite  tomentose  to  subshining; 
legs  of  females  dark;  haustellum  length 
varied;  scutellum  lighter  in  color  than 
rest  of  notum  in  most  individuals 17 

17a.  Parafacial  hairs  pale  and,  in  most  indi- 
viduals, small  (Fig.  97)  (west  only  to  Ar- 
izona and  Rocky  Mountains) 18 

17b.  Parafacial  hairs  light  brown  to  black, 
short  or  medium  in  length  (Pacific  coast 
states) 19 

18a.  Infrasquamal  setulae  absent  or  greatly 
reduced  in  number  in  most  individuals; 
parafacial  hairs  only  on  upper  anterior 
parafacial  (Fig.  102);  length  of  palpi 
equal  to  or  slightly  less  than  half  length 
of  haustellum,  broadened  at  tip  (central 
and  eastern  U.S.)....  incerta  (West)  (in  part) 

18b.  Infrasquamal  setulae  present  in  most  in- 
dividuals; parafacial  hairs  extending 
ventrally  on  face  to  level  of  apex  of  an- 
tennae (Fig.  97);  length  of  palpi  greater 
than  half  length  of  haustellum,  narrow 

(southwestern  U.S.  into  Texas)  

major  (Bigot) 

19a(17b).  Third  antennal  segment  subequal  to 


or  shorter  than  second  segment;  smallest 
distance  between  eyes  of  male  subequal 
to  width  of  frontal  vitta  at  antennal  base; 
female  with  parafacial  hairs  confined  to 
area  near  antennae;  facial  tomentum  dull 
(southern  California)  sabroskyi  n.sp. 

19b.  Third  antennal  segment  longer  than  sec- 
ond segment;  smallest  distance  between 
eyes  of  male  less  than  width  of  frontal 
vitta  at  base  of  antennae;  females  with 
parafacial  hairs  scattered,  often  along 
center  of  parafacial;  facial  tomentum 

shining  (throughout  California)  

calif  arnica  n.sp. 

20a.  Haustellum  length  greater  than  0.6  times 
head  height,  thin,  narrowed  apically  21 

20b.  Haustellum  length  less  than  or  equal  to 
0.6  times  head  height,  broad  and  linear 
in  most  individuals  24 

2 la.  Length  of  plumosity  on  arista  more  than 
twice  length  of  second  antennal  segment; 
haustellum  length  0.65  to  0.7  times  head 
height,  narrow  (Fig.  81)  (northern  and 
eastern  U.S.)  harpasa  (Walker) 

21b.  Length  of  plumosity  on  arista  less  than 
or  equal  to  twice  length  of  second  anten- 
nal segment;  haustellum  length  varied 
(western  U.S.)  22 

22a.  Parafacial  hairs  long,  dark,  and  abun- 
dant, uniformly  covering  parafacial  (Fig. 
26);  haustellum  length  0.6  to  0.8  times 
head  height,  narrowed  apically  in  most 
individuals;  abdomen  reddish  laterally  __ 
conjuncta  (Wulp) 

22b.  Parafacial  hairs  sparse  (Fig.  41);  haus- 
tellum  length   and   abdomen   varied 
23 

23a.  Length  of  haustellum  more  than  0.7 
times  head  height,  strongly  narrowed 
apically;  parafacial  hairs  sparse,  located 
only  on  anterior  portion  of  parafacial 
(Fig.  41);  male  abdomen  dark  brown  or 
gray  with  little  if  any  reddish  color;  ninth 
tergite  with  several  strong  setae  (Fig.  38) 

(Texas  and  Oklahoma)  

westi  n.sp.  (in  part) 

23b.  Length  of  haustellum  less  than  0.65 
times  head  height,  broad;  not  narrowed 
apically  in  most  individuals;  parafacial 
hairs  occurring  along  center  of  parafacial 
(Fig.  46);  male  abdomen  distinctly  red- 
dish with  longitudinal  stripe;  ninth  ter- 


WILDER:  NEARCTIC  PTILODEXIA 


19 


gite  with  hairs  only  (Pacific  coast  states) 
calif ornica  n.sp.  (in  part) 

24a(20b).  Abdomen  blackish  with  little  if  any 
orange  coloration  laterally;  parafacial 
hairs  inserted  close  to  anterior  margin  of 
eye  in  most  individuals;  palpi  very  short, 
less  than  half  length  of  haustellum  (Fig. 
21);  wings  of  many  males  basally  dark- 
ened    obscura  West 

24b.  Abdomen,  especially  of  male,  with  at 
least  some  red  or  orange  coloration  lat- 
erally; parafacial  hairs  not  inserted  close 
to  anterior  eye  margin  in  most  individu- 
als; palpi  longer  than  half  length  of  haus- 
tellum (Fig.  51);  wings  of  males  not  ba- 
sally darkened  25 

25a.  Femora  of  members  of  both  sexes  or- 
ange or  marked  with  orange;  abdominal 

tomentum  concolorous  (California) 

pacifica  n.sp. 

25b.  Femora  of  members  of  both  sexes  brown 
or  black  (some  specimens  from  Texas 
may  have  orange  femora);  abdominal  to- 
mentum bicolored  (widespread) 26 

26a.  Epistomal  angle  of  head  not  prominent 
(Fig.  61);  females  with  6  to  8  dorsal  and 
lateral  marginal  setae  on  abdominal  seg- 
ment III;  males  with  1  pair  of  dorsal  mar- 
ginal setae  on  abdominal  segment  II; 
haustellum  length  0.4  to  0.5  times  head 

height  (central  and  eastern  U.S.) 

carolinensis  Brauer  and  Bergenstamm 

(in  part) 

26b.  Epistomal  angle  of  head  prominent  (Fig. 
46);  females  with  10  to  12  dorsal  and  lat- 
eral marginal  setae  on  abdominal  seg- 
ment III;  most  males  with  2  pairs  of  dor- 
sal marginal  setae  on  abdominal  segment 
II;  haustellum  length  0.5  to  0.65  times 

head  height  (western  U.S.) 

calif  ornica  n.sp.  (in  part) 

27a(lb).  Abdomen  without  any  reddish  or  or- 
ange coloration  laterally  in  most  males; 
tip  of  abdomen  and  genitalia  generally 
reddish  yellow;  facial  tomentum  strongly 
shining;  length  of  plumosity  on  arista 
more  than  twice  length  of  second  anten- 
nal  segment;  face  appearing  narrow  (Fig. 
87);  infrasquamal  setulae  absent  in  many 
individuals;  femora  of  many  males  yel- 
low (central  and  eastern  U.S.  into  New 


Mexico  and  British  Columbia) 

rufipennis  (Macquart) 

27b.  Abdomen  orange  laterally  in  most  males; 
tip  of  abdomen  not  noticeably  lighter 
than  rest  of  abdomen;  facial  tomentum 
rather  dull;  length  of  plumosity  on  arista 
at  most  equal  to  twice  length  of  second 
antennal  segment;  face  appearing  broad 
(Fig.  92);  infrasquamal  setulae  present; 
femora  of  males  dark  (Utah  and  southern 
Idaho  south  into  Mexico)  arida  (Curran) 

The  Nearctic  Species  of  Ptilodexia 

agilis  Group 
Ptilodexia  agilis  Reinhard 

(Figures  8-12) 

Ptilodexia  agilis  REINHARD,  1943:22.  SABROSKY  AND  AR- 
NAUD  (1965:988).  [HOLOTYPE,  male,  deposited  in  CNCI, 
labeled,  College  Station,  Texas,  8  Oct.  1933,  H.  J.  Rein- 
hard.] 

TAXONOMIC  NOTES. — Although  I  have  not 
seen  the  holotype  of  P.  agilis,  members  of  this 
species  are  quite  distinctive,  and  the  original 
description  is  adequate  to  assure  the  identity  of 
the  specimens  examined.  Even  though  P.  agilis 
was  described  from  a  disjunct  population,  the 
type-specimens  are  typical  of  the  species.  Type 
material  is  reported  to  be  in  excellent  condition. 

DIAGNOSIS. — Ptilodexia  agilis  is  a  distinct 
species,  members  of  which  can  be  separated 
from  their  congeners  by  the  following  character 
states:  size  small;  parafacial  hairs  fine,  medium 
in  length,  inserted  below  apex  of  antennae  or 
less  than  0.25  mm  from  anterior  margin  of  eye 
in  only  a  few  individuals;  haustellum  less  than 
half  head  height;  length  of  plumosity  on  the  aris- 
ta 1.5  to  2.0  times  length  of  second  antennal 
segment;  mediotergite  polished;  abdomen  and 
scutellum  entirely  blackish;  female  with  legs 
pale  colored. 

MATERIAL  EXAMINED. — Specimens  exam- 
ined include  812  males  and  488  females,  data  as 
listed  by  Wilder  (1976). 

DISTRIBUTION. — Ptilodexia  agilis  ranges  from 
Arizona  and  New  Mexico  north  through  Cali- 
fornia, the  Great  Basin,  and  the  Rocky  Moun- 
tains into  Alberta  and  British  Columbia.  There 
are  a  few  records  from  eastern  Texas,  where  the 
topotypic  population  is  found. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  April  to  October,  and  adults  can  be  col- 


20 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


10 


FIGURES  8-12.    Ptilodexia  agilis  Reinhard.  Fig.  8.  Genitalia  of  male,  posterior  view.  Fig.  9.  Ejaculatory  apodeme.  Fig.  10. 
Genitalia  of  male,  lateral  view.  Fig.  11.  Head  of  male,  lateral  view.  Fig.  12.  Distribution  of  P.  agilis. 


lected  at  any  time  during  this  period.  July  and 
August  are  the  most  frequent  months  of  collec- 
tion, but  local  variation  is  common.  In  Califor- 
nia, for  example,  P.  agilis  adults  are  collected 
more  frequently  in  September  and  October 
along  the  coast  and  in  the  south;  but  in  the  Sier- 
ra, northern  California,  and  Oregon,  June  and 
July  are  the  main  periods  of  activity. 

This  species  inhabits  both  mountains  and  low- 
lands. Adults  have  been  collected  at  elevations 
up  to  2,600  m  in  Arizona,  3,800  m  in  California, 
above  the  3,000-m  level  in  Colorado,  and  fre- 
quently above  3,000  m  elsewhere.  Ptilodexia 
agilis  adults  have  frequently  been  taken  at  low 
elevations  in  such  areas  as  the  San  Joaquin  Val- 
ley, the  Great  Basin,  and  eastern  Texas.  Spec- 
imens have  been  collected  by  UV  light  trap, 
Malaise  trap,  wind  vane  trap,  and  by  sweeping 
foliage. 

Flowers  visited  include  the  Compositae  Achil- 
lea  Millefolium,  Chrysothamnus  viscidiflorus , 
Baccharis  glutinosa,  B.  pilularis,  Eriogonum 
nudum,  Solidago  trinervata,  and  Lepidospartum 


squamatum.  Unidentified  Compositae  visited 
were  Achillea  sp.,  Baccharis  sp.,  Solidago  sp., 
Eriogonum  sp.,  and  Haplopappus  sp.  Other 
flowers  from  which  P.  agilis  adults  have  been 
collected  are  Allium  (Liliaceae)  and  Salix  (Sal- 
icaceae).  At  two  localities  in  Colorado,  speci- 
mens were  collected  on  Dwarf  Mistletoe  (Ar- 
ceuthobium  cyanocarpum)',  and  members  of  this 
species  are  believed  to  be  pollinators  of  that 
plant.  One  specimen  was  collected  in  an  emer- 
gence trap  under  a  filbert  tree  in  Oregon. 

There  are  no  data  on  the  life  history  of  this 
insect.  Its  members  are  probably  not  host  spe- 
cific, judging  from  the  diverse  assortment  of 
habitats  and  wide  geographical  and  temporal 
ranges  of  the  species. 

Ptilodexia  mathesoni  (Curran) 

(Figures  13-17) 

Rhynchiodexia  mathesoni  CURRAN,  1931:93.  WEST  (1950:110); 
SABROSKY  AND  ARNAUD  (1965:989).  [LECTOTYPE  (here  des- 
ignated), male,  deposited  in  CUIC,  labeled,  "Douglas  Lake, 
Mich.,  24-VII-22"/"Wing  Slide,  Cornell  U.,  Lot.  919,  Sub. 
138,  L.  S.  West"/"cJ  Holotype  Rhynchiodexia  mathesoni 


WILDER:  NEARCTIC  PTILODEXIA 


21 


16 


FIGURES  13-17.    Ptilodexia  mathesoni  (Curran).  Fig.  13.  Genitalia  of  male,  posterior  view.  Fig.  14.  Ejaculatory  apodeme. 
Fig.    15.  Genitalia  of  male,  lateral  view.  Fig.  16.  Head  of  male,  lateral  view.  Fig.  17.  Distribution  of  P.  mathesoni. 


Curran'T'Holotype  Cornell  U.  No.  1938"/"Cornell  U.  Lot. 
922,  Sub.  40"/"Lectotype  Rhynchiodexia  mathesoni  Cur- 
ran  designated  by  D.  Wilder,  1976."] 

TAXONOMIC  NOTES. — Even  though  the  label 
on  the  type-specimen  reads  "Holotype  Rhyn- 
chiodexia mathesoni  Curran,"  this  specimen  is 
not  a  holotype.  The  author  of  the  paper  vali- 
dating the  species  made  no  mention  of  type  ma- 
terial or  of  type-locality.  The  type  label  with 
Curran  as  author  was  put  on  the  specimen  at  a 
later  date;  it  was  West's  "holotype,"  not  Cur- 
ran's.  The  name  P.  mathesoni  was  proposed  by 
West  and  validated  by  Curran  in  his  1931  key. 
I  have  designated  West's  "holotype"  as  the  lec- 
totype.  It  is  a  large  specimen  in  excellent  con- 
dition. 

There  is  one  other  specimen  which  I  believe 
Curran  had  before  him  while  writing  his  key. 
This  specimen,  also  from  Douglas  Lake,  Mich- 
igan, is  deposited  in  AMNH  and  was  collected 
on  the  same  date  as  the  specimen  West  labeled 
as  holotype.  I  have  designated  this  specimen  as 
a  paralectotype.  It  might  be  argued  that  this  sec- 


ond specimen,  because  it  is  labeled  with  Curran 
as  the  author,  was  later  sent  to  AMNH  and  not 
seen  by  Curran.  However,  the  type  labels  of  all 
five  of  the  species  validated  in  Curran's  key  give 
Curran  as  the  author.  I  believe  that  all  these 
labels  were  changed  at  a  later  date,  and  since 
Curran  makes  no  mention  of  material,  I  feel  that 
these  two  identically  labeled  specimens  are  ac- 
tually syntypes. 

DIAGNOSIS. — Ptilodexia  mathesoni  is  a  dis- 
tinctive species,  evidenced  by  the  following 
combination  of  character  states:  body  color 
black,  shining;  parafacial  hairs  long,  black,  in- 
serted ventral  to  apex  of  antennae  or  less  than 
0.25  mm  from  anterior  margin  of  eye  in  only  a 
few  individuals;  haustellum  less  than  half  head 
height  in  length;  width  of  arista  and  plumosity 
1.5  to  2  times  length  of  second  antennal  seg- 
ment; mediotergite  polished;  wings  dark  in  col- 
or; male  abdomen  with  pollen  evenly  distribut- 
ed; female  with  pale-colored  legs. 

MATERIAL  EXAMINED. — Twenty-one  males 
and  eleven  females  were  examined. 


22 


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21 


FIGURES  18-22.    Ptilodexia  obscura  West.   Fig.  18.  Genitalia  of  male,  posterior  view.   Fig.  19.  Ejaculatory  apodeme.   Fig. 
20.  Genitalia  of  male,  lateral  view.  Fig.  21.  Head  of  male,  lateral  view.   Fig.  22.  Geographical  distribution. 


DISTRIBUTION. — The  species,  although  rec- 
ords are  few,  seems  to  range  through  the  north- 
eastern United  States  and  eastern  Canada. 
There  is  one  record  from  Victoria  Beach,  Man- 
itoba. 

BIOLOGICAL  NOTES. — The  flight  period  is 
from  April  to  October  with  the  main  period  of 
activity  between  mid-June  and  mid-August.  All 
records  for  months  other  than  July  and  August 
are  from  Suffolk  County,  New  York. 

There  are  no  life-history  data  for  P.  mathe- 
soni.  One  male  was  collected  on  Solidago  can- 
adensis. 

Ptilodexia  obscura  West 

(Figures  18-22) 

Ptilodexia  obscura  WEST,  1925:133.  LEONARD  (1928:822); 
CURRAN  (1930:93);  SABROSKY  AND  ARNAUD  (1965:989). 
[HoLOTYPE,  female,  deposited  in  SIIS,  labeled,  "Wading 
River,  L.I.,  June  29,  1917,  W.  T.  Davis."] 

TAXONOMIC  NOTES. — West  described  P.  ob- 
scura from  three  female  specimens,  one  of 
which  he  designated  holotype.  The  two  para- 
types  are  so  labeled  and  are  deposited  in  CUIC. 


They  closely  resemble  the  holotype.  There  is  a 
male  specimen  from  Victoria  Beach,  Manitoba, 
deposited  in  AMNH,  which  bears  a  handwritten 
label  reading,  "/?.  obscura  West."  It  appears 
that  West  recognized  the  male  of  the  species  at 
a  later  time,  even  though  this  particular  male 
specimen  differs  considerably  from  the  female 
type-series.  West  still  used  the  generic  name 
Ptilodexia  for  obscura  in  1950,  but  because  his 
concept  of  the  genus  changed,  it  is  possible  that 
he  might  have  written  "/?.  obscura"  instead  of 
"P.  obscura."  It  is  doubtful  that  Curran  labeled 
the  specimen  or  ever  looked  at  West's  types, 
since  in  his  1931  key,  he  describes  P.  obscura 
specimens  as  being  over  12.5  mm  long. 

DIAGNOSIS. — Ptilodexia  obscura  is  a  fairly 
distinctive  species.  Its  members  may  be  sepa- 
rated from  their  congeners  by  the  following 
combination  of  character  states:  face  narrow, 
with  parafacial  hairs  abundant,  long,  fine,  dark, 
and  inserted  below  lower  edge  of  eye  and  close 
to  its  anterior  edge;  third  antennal  segment  dark 
in  most  specimens,  broadened  apically;  length 
of  plumosity  on  arista  1.5  times  length  of  second 


WILDER:  NEARCTIC  PT1LODEXIA 


26 


FIGURES  23-27.    Ptilodexia  conjuncta  (Wulp).    Fig.  23.  Genitalia  of  male,  posterior  view.   Fig.  24.  Ejaculatory  apodeme. 
Fig.  25.  Genitalia  of  male,  lateral  view.   Fig.  26.  Head  of  male,  lateral  view.   Fig.  27.  Geographical  distribution. 


segment;  haustellum  broad,  length  0.55  to  0.6 
times  head  height;  palpi  short,  half  haustellum 
length  or  less;  notum  covered  with  short,  ap- 
pressed,  grayish  pollen,  giving  it  a  smooth,  sub- 
shining  appearance;  legs  of  females  brown;  ab- 
dominal color,  grossly  appearing  black,  but 
actually  rufescent  laterally. 

MATERIAL  EXAMINED. — One  hundred  forty- 
two  males  and  53  females  of  P.  obscura  were 
examined. 

DISTRIBUTION. — The  range  of  P.  obscura  ex- 
tends from  New  Brunswick  south  into  the  Geor- 
gia Appalachians  west  into  Arkansas,  eastern 
Kansas  and  Nebraska,  western  South  Dakota 
and  Saskatchewan,  and  Manitoba. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  April  to  September  with  June  and  July  the 
most  common  months  of  collection,  a  later  av- 
erage seen  only  in  Manitoba.  Most  specimens 
have  been  collected  at  low  elevations,  the  ex- 
ceptions coming  from  800-1,700  m  in  the  Ap- 
palachians in  Tennessee,  North  Carolina,  and 
Virginia. 

Collecting  methods  yielding  specimens  of  P. 
obscura  include  sweeping  and  Malaise  trap. 


Specimens  have  been  collected  on  the  flowers 
of  Solidago  canadensis  (Compositae),  and 
Aruncus  sp.  and  Spiraea  latifolia,  both  Rosa- 
ceae. 

conjuncta  Group 
Ptilodexia  conjuncta  (Wulp) 

(Figures  23-27) 

Rhynchodexia  conjuncta  WULP,  1891:228.  ALDRICH  (1905:499); 
GUIMARAES  (1971:33).  [LECTOTYPE  (here  designated), 
male,  deposited  in  BMNH,  labeled  "Lectotype"/"tJ'V 
"B.C. A.  Dipt.  II,  Rhynchodexia  conjuncta  v.d.W. "/"Cen- 
tral America.  Pres.  by  F.  D.  Godman.  O.  Salvin.  1903- 
172"/"Ciudad,  Mex.,  8100  ft.,  Forrer'VLectotype  Rhyn- 
chodexia  conjuncta  Wulp  designated  by  D.  Wilder,  1975."] 

Rhynchodexia  simulans  WULP,  1891:229.  ALDRJCH  (1905:499); 
GUIMARAES  (1971:34).  [LECTOTYPE  (here  designated), 
male,  deposited  in  BMNH,  labeled,  "Lectotype'V'N.  So- 
nora,  Mexico.  Morrison"/"  d"/"B.  C.  A.  Dipt.  II,  Rhyn- 
chodexia simulans,  v.d.W. "/"Central  America,  pres.  by  F. 
D.  Godman,  O.  Salvin.  1903-172"/"Lectotype  Rhyncho- 
dexia simulans  Wulp,  designated  by  D.  Wilder,  1975."]  NEW 
SYNONYMY 

Ptilodexia  tibialis  (partim):  ALDRICH  (1905:504). 

TAXONOMIC  NOTES. — Wulp  described  P. 
conjuncta  from  two  male  cotypes,  one  of  which 
I  have  designated  as  lectotype.  This  specimen 


24 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


is  in  good  condition  except  for  a  few  broken 
setae. 

Wulp  described  P.  simulans  and  P.  conjuncta 
in  the  same  paper.  The  cotype  (one  of  two) 
which  I  saw  and  designated  lectotype  is  in  good 
condition,  although  it  has  the  dorsal  setae  and 
abdomen  broken. 

Wulp  realized  that  P.  conjuncta  and  P.  sim- 
ulans were  very  closely  related.  He  separated 
members  of  each  on  the  basis  of  size,  curvature 
of  the  hind  tibia,  and  some  other  minor  char- 
acters. Size,  of  course,  cannot  be  used  effec- 
tively to  separate  these  parasitic  flies.  A  curved 
hind  tibia  is  a  character  state  which  occurs  fre- 
quently in  members  of  many  species  of  Ptilo- 
dexia,  especially  in  those  of  P.  conjuncta,  where 
it  constitutes  part  of  normal  intraspecific  varia- 
tion. 

DIAGNOSIS. — Ptilodexia  conjuncta  is  a  vari- 
able species.  Its  members  can  be  distinguished 
by  the  following:  face  wide;  parafacial  hairs 
long,  fine,  dark,  abundant,  inserted  below  level 
of  lower  edge  of  eye  in  some  specimens  and  at 
least  0.12  mm  from  anterior  edge  of  eye  in  all 
but  a  few  specimens;  haustellum  long,  ranging 
from  0.6  to  0.8  times  head  height,  slender,  nar- 
rowed apically  in  most  specimens;  carina  fairly 
well  developed;  width  of  arista  and  plumosity 
1.5  to  2  times  length  of  second  antennal  seg- 
ment; oral  margin  distinctly  protruding;  scutel- 
lum  and  sides  of  abdomen  reddish. 

MATERIAL  EXAMINED. — Specimens  exam- 
ined included  1,238  males  and  629  females. 

DISTRIBUTION. — This  species  ranges  from 
British  Columbia  and  Alberta  south  through  the 
Rocky  Mountains  into  Mexico.  There  are  scat- 
tered records  from  the  plains  states,  the  Great 
Basin,  California,  and  Oregon.  There  are  a  few 
doubtful  records  from  the  eastern  United  States. 

BIOLOGICAL  NOTES. — The  flight  period  ranges 
from  February  to  October.  Most  of  the  activity 
occurs  in  July  and  August,  earlier  in  the  north- 
ern areas  and  later  further  south.  In  Arizona 
(and  possibly  Texas),  P.  conjuncta  appears  to 
have  two  broods,  one  in  March  and  April,  and 
another  in  August  and  September. 

Ptilodexia  conjuncta  adults  are  generally 
found  in  mountainous  areas  at  altitudes  from 
1,500  to  2,750  m  and  in  some  areas  up  to  3,660 
m.  They  are  also,  although  less  commonly,  col- 
lected at  low  elevations  in  coastal  as  well  as  in- 
land areas.  Label  data  indicate  that  they  have 
been  collected  in  meadow  sweeps,  in  a  meadow 


in  spruce-fir  zone  (2,750  m),  in  pine-spruce-as- 
pen zone,  and  above  timberline.  Productive  col- 
lecting methods  for  P.  conjuncta  specimens  are 
UV  light  and  Malaise  trap. 

Specimens  of  P.  conjuncta  have  been  col- 
lected from  many  flowers.  These  include:  Se- 
necio  salignus,  Heliopsis  parvifolia,  Cacalia  de- 
composita,  Enceliafarinosa,  Geraea  canescens, 
Chrysothamnus  greeni,  Achillea  Millefolium; 
and  unidentified  species  of  Ence I'm  sp.,  Gutier- 
rezia  sp.,  Solidago  sp.,  Baccharis  sp.,  Helian- 
thus  sp.,  Senecio  sp.,  Eriogonum  sp.,  Achillea 
sp.,  Bigelovia  sp.,  Aster  sp.,  and  Rudbeckia  sp. 
(all  Compositae).  Other  flowers  visited  include, 
Ceanothus  fendleri  and  unidentified  Ceanothus 
sp.  (Rhamnaceae);  Arctostaphylos  sp.  (Erica- 
ceae); Melilotus  sp.,  and  Dale  a  sp.  (Fabaceae); 
Lippia  wrightii  (Verbenaceae);  and  Arceuthob- 
ium  sp.  (Loranthaceae).  Ptilodexia  conjuncta 
adults,  along  with  those  of  P.  agilis,  have  been 
observed  pollinating  Dwarf  Mistletoe  (Arceu- 
thobium  cyanocarpum). 

Ptilodexia  planifrons  (Wulp) 

(Figures  28-32) 

Rhynchodexia  planifrons  WULP,  1891:234.  ALDRICH  (1905:499); 
GUIMARAES  (1971:33).  [HOLOTYPE,  male,  deposited  in 
BMNH,  labeled,  "Holotype'VCiudad,  Mexico,  8100  ft., 
Ferrer"/"  d"/"B.  C.  A.  Dipt.  II,  Rhynchodexia  planifrons, 
v.d.W'VCentral  America  pres.  by  F.  D.  Godman,  O.  Sal- 
vin.  1903-172."] 

Dexia  harpasa  (partim):  ALDRICH  (1925:114).  (Misidentifica- 
tion). 

TAXONOMIC  NOTES. — The  holotype  of  this 
species,  deposited  in  BMNH,  is  in  poor  condi- 
tion, but  is  still  recognizable.  The  facial  band  is 
obscured,  the  tibiae  are  quite  light  and  distinctly 
curved  (a  frequently  encountered  anomaly  in 
species  of  Ptilodexia),  and  the  striping  on  the 
notum  is  more  distinct. 

Labels  on  the  specimen  say  only  "Ciudad, 
Mexico,  8100  ft.,"  but  in  the  description,  the 
origin  of  this  specimen  is  stated  as  "Ciudad  in 
Durango,  8100  ft."  I  follow  Wulp's  original  pub- 
lication in  calling  Durango  the  type-locality. 

DIAGNOSIS. — Ptilodexia  planifrons  is  a  dis- 
tinctive species  and  its  members  can  be  identi- 
fied by  the  following  character  combination: 
face  wide;  presence  of  a  contrasting  tomentose 
diagonal  band  extending  from  antennal  base  to 
eye  margin;  parafacial  hairs  dark,  coarse,  abun- 
dant, inserted  lower  than  level  of  oral  vibrissae 
only  in  a  few  specimens;  antenna  with  plumosity 


WILDER:  NEARCTIC  PTILODEXIA 


25 


28 


31 


FIGURES  28-32.    Ptilodexia  planifrons  (Wulp).   Fig.  28.  Genitalia  of  male,  posterior  view.    Fig.  29.  Ejaculatory  apodeme. 
Fig.  30.  Genitalia  of  male,  lateral  view.   Fig.  31.  Head  of  male,  lateral  view.  Fig.  32.  Geographical  distribution. 


on  arista  less  than  or  equal  to  length  of  second 
antennal  segment;  carina  long,  slightly  keeled 
and  prominent;  two  pairs  of  oral  vibrissae  in 
most  specimens;  frontal  vitta  very  wide  at  an- 
tennal base;  haustellum  long,  0.7  to  0.85  times 
head  height,  narrowed  apically;  scutellum  and 
abdomen  entirely  dark  gray  or  black. 

MATERIAL  EXAMINED. — One  hundred  sev- 
enty-three males  and  70  females  of  P.  planifrons 
were  examined. 

DISTRIBUTION. — This  species  ranges  from 
Flagstaff  south  through  the  mountains  of  central 
and  southeastern  Arizona,  through  the  central 
mountainous  region  of  New  Mexico,  the  west- 
ern tip  of  Texas,  and  south  into  Durango,  Mex- 
ico. There  is  one  record  from  Colorado;  how- 
ever, no  exact  locality  is  given. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  August  through  October,  with  the  majority 
of  records  from  mid-August  to  mid-September. 
The  earliest  seasonal  record  is  one  specimen 
collected  on  27  June  (error?)  from  El  Paso,  Tex- 
as, in  1921,  and  the  latest  is  a  series  of  24  females 
collected  on  22  October  1964,  in  Cochise  Coun- 
ty, Arizona. 


Ptilodexia  planifrons  adults  generally  occur  in 
the  mountains  and  have  been  collected  at  many 
elevations  between  1,370  and  3,350  m.  Flowers 
visited  include  Gutierrezia  sarothrae  (1,800- 
2,440  m,  Apache  County,  Arizona),  Heliopsis 
parvifolia  (2,590  m,  Chiricahua  Mts.,  Cochise 
County,  Arizona),  Solidago  trinervata  (Sierra 
Madre,  2,230  m),  and  Rudbeckia  sp.  in  Chihua- 
hua, all  Compositae.  All  except  two  of  the 
flower-visiting  flies  were  males  (one  female  col- 
lected on  Heliopsis  sp.  and  another  at  Rudbeck- 
ia sp.).  One  specimen  was  collected  at  a  light. 

Ptilodexia  contristans  (Wulp) 

(Figures  33-37) 

Hystrichodexia  contristans  WULP,  1891:221.  [HOLOTYPE, 
male,  deposited  in  BMNH,  labeled,  "Holotype'VOmil- 
teme,  Guerrero,  8000  ft.,  July,  H.  H.  Smith"/"  d"/"B.  C. 
A.  Dipt.  II.,  Hystrichodexia  contristans,  v.d.W. "/"Central 
America,  Pres.  by  F.  Godman,  O.  Salvin.  1903-172."] 

Rhynchodexia  punctipennis  WULP,  1891:233.  ALDRICH 
(1905:499);  GUIMARAES  (1971:33).  [LECTOTYPE  (here  des- 
ignated), male,  deposited  in  BMNH,  labeled,  "Cotype"/ 
"Sierra  de  las  Aguas  Escondidas,  Guerrero,  9500  ft.,  July. 
H.  H.  Smith"/"  <J"/"B.  C.  A.  Dipt.  II.  Rhynchodexia  punc- 
tipennis, v.d.W. "/"Central  America.  Pres.  by  F.  D.  God- 
man, O.  Salvin.  1903-172"/"Lectotype  Rhynchodexia 


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36 


FIGURES  33-37.    Ptilodexia  contristans  (Wulp).  Fig.  33.  Genitalia  of  male,  posterior  view.  Fig.  34.  Genitalia  of  male,  lateral 
view.  Fig.  35.  Ejaculatory  apodeme.  Fig.  36.  Head  of  male,  lateral  view.  Fig.  37.  Geographical  distribution. 


punctipennis  Wulp  designated  by  D.  Wilder  1975."]  NEW 
SYNONYMY. 

Ptilodexia  constrictans  (Wulp):  GUIMARAES  (1971:33)  [lapsus 
calamus]. 

TAXONOMIC  NOTES. — The  holotype  of  this 
species  is  in  fair  condition  except  for  a  broken 
thorax  and  an  abdomen  which  is  oily,  obscuring 
the  tomentum  patterns. 

Wulp  described  Rhynchodexia  punctipennis 
and  Hystrichodexia  contristans  in  the  same  pa- 
per. He  felt  that  the  two  genera  differed  by  two 
superficial  characters,  the  general  body  shape 
and  the  hairs  of  the  abdomen.  The  lectotype  of 
P.  punctipennis,  although  in  poor  condition  is 
certainly  a  teneral  specimen  of  P.  contristans. 

DIAGNOSIS. — Specimens  of  P.  contristans  can 
be  separated  from  those  of  related  species  by 
the  following  combination  of  characters:  face 
very  wide;  parafacial  hairs  long,  dark,  abundant, 
occurring  to,  and  in  most  specimens,  also  below 
level  of  lower  eye  margin;  third  antennal  seg- 
ment broadened  apically  in  some  individuals; 
plumosity  on  arista  short,  its  width  subequal  to 


length  of  second  antennal  segment;  carina  short 
but  well  developed;  haustellum  narrow,  equal  to 
0.6  times  head  height;  oral  margin  strongly  pro- 
truding; scutellum  and  sides  of  abdomen  reddish 
laterally,  wing  base  distinctly  darkened;  ninth 
tergite  with  several  strong  setae. 

MATERIAL  EXAMINED. — Ninety-one  males 
and  20  females  were  examined. 

DISTRIBUTION. — This  species  ranges  from  the 
central  and  southeastern  mountains  of  Arizona 
southward  into  the  mountains  in  Durango,  Ve- 
racruz, and  Guerrero.  There  is  one  record  from 
New  Mexico. 

BIOLOGICAL  NOTES. — The  flight  period  is 
from  mid- August  to  early  September,  with  a  few 
scattered  records  in  July.  The  holotype  was  col- 
lected in  July. 

All  specimens  were  collected  in  mountainous 
areas.  The  lowest  elevation  indicated  on  labels 
is  2,130  m,  in  Veracruz,  the  highest,  2,900  m,  in 
Guerrero.  Specimens  have  been  collected  on  the 
following  Compositae:  He liopsis  parvifolia ,  Ca- 
calia  decomposita,  Verbesina  encelioides,  He- 


WILDER:  NEARCTIC  PT1LODEXIA 


27 


FIGURES  38-42.    Ptilodexia  westi,  n.sp.   Fig.  38.  Genitalia  of  male,  posterior  view.   Fig.  39.  Ejaculatory  apodeme.   Fig.  40. 
Genitalia  of  male,  lateral  view.  Fig.  41.  Head  of  male,  lateral  view.   Fig.  42.  Geographical  distribution. 


lenium  hoopesii,  and  unidentified  Senecio  sp., 
Cirsium  sp.,  and  Solidago  sp.  There  are  no  as- 
sociated host  data. 

Ptilodexia  westi,  new  species 

(Figures  38-42) 

TYPE-LOCALITY. — The  holotype  was  collected  at  Imperial, 
Texas,  10  April  1954  by  L.  D.  Beamer. 

TYPE-SPECIMENS. — The  male  holotype  is  deposited  in 
CNCI,  the  allotype,  from  Las  Cruces,  New  Mexico,  in 
USNM.  Complete  data  from  these  specimens  and  the  21  para- 
types  are  listed  below. 

DIAGNOSIS. — This  is  a  distinctive  species,  its 
members  easily  separable  from  those  of  other 
species  in  the  genus  by  the  following  combina- 
tion of  character  states:  face  wide,  tomentum 
grayish,  very  heavy;  parafacial  hairs  from  short 
to  medium  in  length,  occurring  only  on  anterior 
half  of  parafacial,  below  apex  of  antennae  in  a 
few  individuals;  length  of  haustellum  0.7  to  0.8 
times  head  height,  narrowed  apically,  length  of 
plumosity  on  arista  varied,  from  1  to  1.25  times 
length  of  second  antennal  segment;  abdomen 


heavily  tomentose,  in  males  integument  dark 
gray  with  no  median  stripe,  in  females  brown  or 
rufous;  ninth  tergite  of  males  with  several  strong 
setae;  genitalia  unlike  those  of  adults  of  any  oth- 
er species. 

DESCRIPTION. — Male:  Total  body  length  10.5 
mm,  grayish.  Head  height  2.6  mm;  profrons  0.25 
of  head  length.  Facial  tomentum  very  heavy, 
dull,  grayish  white,  obscuring  integumental  col- 
or; parafrontal  hairs  long,  sparse,  dark;  parafa- 
cial hairs  medium  in  length,  dark,  rather  sparse, 
located  along  anterior  half  of  parafacial  ventrally 
to  slightly  beyond  apex  of  antennae.  Width  of 
frontal  vitta  at  antennal  base  0.4  mm,  at  narrow- 
est part,  0.1  mm;  carina  strongly  developed, 
slightly  keeled,  broad,  not  depressed  below  apex 
of  antennae;  facial  cavities  strongly  darkened. 
Height  of  gena  0.25  times  head  height.  One  pair 
of  strong  ocellar  setae,  one  pair  of  smaller  post- 
ocellars  and  postverticals;  one  pair  of  strong  and 
one  pair  of  weak  inner  verticals;  outer  verticals 
subequal  to  postorbitals.  Fifteen  pairs  of  medi- 
um-length postorbital  setae,  some  a  little  shorter 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


than  others;  ventral  to  postorbitals  is  an  irreg- 
ular row  of  dark  hairs  merging  into  dense  whitish 
occipital  hair.  Seven  pairs  of  frontal  setae;  one 
pair  of  oral  vibrissae  0.3  mm  from  oral  margin, 
the  shallow  depression  between  them  0.4  mm 
wide;  9  additional  oral  setae,  one  above  vibris- 
sae quite  strong,  peristomals  gradually  increas- 
ing in  length  with  distance  from  vibrissae.  Epi- 
stome  not  protruding,  its  width  twice  length  of 
first  two  antennal  segments.  Haustellum  narrow, 
length  0.7  times  head  height,  narrowed  apically; 
palpi  testaceous,  narrow,  length  0.5  times  haus- 
tellum  length.  Antennae  dark  testaceous,  third 
segment  brownish  apically,  arista  and  plumosity 
dark  brown.  Third  segment  of  antenna  1.5  times 
length  of  second  antennal  segment.  Thorax: 
Mesonotum  dark  brown  with  heavy  gray  tomen- 
tum;  striping  obscure.  Integument  of  scutellum 
and  postalar  calli  concolorous  with  rest  of  no- 
tum.  Six  humeral  setae,  2  posthumerals  and  1 
presutural,  3  pairs  of  discal  scutellars.  Pleura 
brown,  sutures  lighter,  tomentum  grayish;  infra- 
squamal  setulae  present;  squamae  whitish, 
wings  light  brown;  epaulet  light  brown,  basi- 
costa  testaceous;  mediotergite  tomentose.  Legs 
dark  reddish  brown,  tibiae  lighter  apically.  An- 
terior femur  with  7  dorsal  setae,  8  posterodor- 
sals,  13  posteroventrals;  tibia  with  slightly  dark- 
ened dorsal  line.  Midlle  femur  with  5  short 
anterior  setae  in  2  rows.  Posterior  femur  with  8 
anterodorsals,  5  anteroventrals,  6  posteroven- 
trals, and  no  anterior  setae.  Femora  with  color 
uniform,  tibiae  darkened  on  basal  third,  tarsi 
black.  Abdomen  dark  brownish  black  with 
heavy  gray  tomentum  which  is  slightly  brownish 
around  bases  of  median  marginal  setae  on  sec- 
ond tergite  and  along  posterior  margins  of  third 
tergite.  First  syntergite  with  1  pair  of  lateral 
marginal  setae.  Second  tergite  with  1  pair  of  me- 
dian discals,  1  pair  of  median  marginals,  and  1 
pair  of  lateral  marginal  setae.  Third  tergite  with 
1  pair  of  median  discal  setae,  and  8  dorsal  and 
lateral  marginal  setae.  Genitalia:  Eighth  tergite 
with  sparse  small  brown  hairs.  Ninth  tergite 
long,  covered  with  long  dense  brown  hairs  and 
many  long,  strong  setae;  lateral  swelling  absent. 
Surstylus  small,  narrow,  lateral  angle  smoothly 
rounded,  internal  depression  absent,  mesal  sur- 
face slightly  convex  with  strong  hairs;  basally 
with  strong  setae,  in  profile,  elongate,  heart- 
shaped,  inclined  strongly  in  posteroventral  di- 
rection, partially  obscuring  cerci.  Cerci  with  lat- 
eral lobes  angular,  only  slightly  differentiated 


from  arms;  arms  wide,  height  more  than  twice 
that  of  lateral  lobes;  arms  extending  ventrally 
well  past  apex  of  surstyli;  in  profile,  smoothly 
but  strongly  incurved  with  no  bulge,  tips  point- 
ed, facing  in  anteroventral  direction.  Ejacula- 
tory  apodeme  with  stem  distinct  from  bulb  and 
about  1.3  times  its  length;  bulb  open  on  part  of 
one  side  and  top.  Female:  Similar  to  male  except 
for  usual  sexual  differences  and  the  following. 
General  color  reddish  brown.  Width  of  frontal 
vitta  at  narrowest  point  0.3  mm,  at  antennal  base 
0.4  mm,  narrowest  point  near  antennal  base, 
strongly  widened  to  vertex.  Height  of  gena  0.25 
times  head  height.  Nine  pairs  of  stout  postorbital 
setae  with  smaller  setae  irregularly  interspersed; 
six  pairs  of  frontal  setae.  Notum  brown  with  to- 
mentum heavy;  six  strong  humeral  setae;  squa- 
mae whitish;  pleura  light  reddish  brown.  Legs 
lighter  in  color  than  those  of  male,  tibiae  simi- 
larly marked.  Anterior  femur  with  10  postero- 
ventrals, 8  posterodorsals,  and  10  dorsal  setae. 
Middle  femur  with  5  strong  anterior  setae  in  2 
rows,  1  anteroventral,  and  5  posteroventrals. 
Posterior  femur  with  6  anterodorsals,  4  postero- 
ventrals, 5  anteroventrals,  and  no  anterior  se- 
tae. Abdomen  brownish,  lighter  laterally,  cov- 
ered with  heavy  grayish  tomentum.  First 
syntergite  with  1  pair  of  lateral  marginal  setae. 
Second  tergite  with  1  pair  of  lateral  marginals 
and  1  pair  of  median  marginal  setae.  Third  ter- 
gite with  1  pair  of  median  discals  and  10  dorsal 
and  lateral  marginals. 

VARIATION. — Ptilodexia  westi  adults  are  quite 
distinct,  but  even  among  the  small  number  of 
specimens  examined,  a  certain  degree  of  varia- 
tion is  seen.  Total  body  length  ranges  from  10 
to  12  mm.  Facial  tomentum  is  yellow  or  gray  in 
a  few  specimens;  parafacial  hairs  do  not  extend 
to  below  the  antennal  apex  in  most  individuals, 
and  the  hairs  are  varied  in  length,  from  short  to 
long.  The  carina  is  shorter  and  not  visible  be- 
yond the  apices  of  the  antennae  in  some  speci- 
mens. There  are  two  rows,  instead  of  one,  of 
dark  hairs  ventral  to  the  postorbitals  in  a  few 
adults.  The  haustellum  length  is  varied,  from  0.7 
to  0.8  times  the  head  height.  The  antennae  are 
also  varied,  from  rufotestaceous  to  light  brown, 
with  the  third  segment  not  darkened. 

The  wings  of  some  specimens  are  light  in  col- 
or. The  femora  are  darkened  apically  on  all  ex- 
cept the  dorsal  surfaces  in  some  individuals;  this 
is  especially  noticeable  in  females,  where 
ground  color  of  the  femora  can  be  pale  in  color. 


WILDER:  NEARCTIC  PTILODEXIA 


29 


FIGURES  43-47.    Ptilodexia  californica,  n.sp.  Fig.  43.  Genitalia  of  male,  posterior  view.  Fig.  44.  Ejaculatory  apodeme.  Fig. 
45.  Genitalia  of  male,  lateral  view.  Fig.  46.  Head  of  male,  lateral  view.   Fig.  47.  Geographical  distribution. 


In  a  few  specimens,  the  abdomen  is  without 
brownish  tomentum,  and  the  integument  is  red- 
dish brown  ventrally  and  laterally.  The  abdomen 
of  many  females  is  reddish  with  a  vague  narrow 
median  stripe,  and  in  most  females,  it  has  no 
median  discal  setae. 

MATERIAL  EXAMINED. — Fifteen  males  and  8  females  were 
examined  from  the  following  localities.  Arizona:  Pima  Co.: 
IcJ,  Brown's  Canyon,  Baboquivari  Mts.,  18  Aug.  1955,  G. 
Butler  (UAIC).  Kansas:  Clark  Co.:  19,  Sitka,  12  June  1960, 
W.  Van  Velzen  (MSUC).  New  Mexico:  Dona  Ana  Co.:  15, 
Las  Cruces,  21  Sep.  (USNM).  Oklahoma:  Woodward  Co.: 
IcJ,  Range  1  mile  NW  Supply,  1  June  1961,  D.  Bryan,  7<3<$- 
399,9  June,  26 63 9  9 ,  15  June  (OSEC).  Texas:  Brazos  Co. : 
Id,  College  Station,  30  May  1936,  student  collector  (T AMU). 
Pecos  Co.:  Id,  Fort  Stockton,  28  Sep.  1935  (TAMU);  IcJ, 
Imperial,  10  Apr.  1954,  L.  Beamer  (CNCI).  Uvalde  Co.:  \<3, 
Uvalde,  18  May  1914,  Bishopp  (USNM). 

Although  few  specimens  were  available  for 
study,  this  species  appears  to  be  the  most  un- 
usual Ptilodexia  in  the  Nearctic  region.  Its 
members  are,  in  fact,  so  dissimilar  from  those 
of  other  species  that  it  may  ultimately  be  as- 
signed to  a  new  genus. 

BIOLOGICAL    NOTES. — The    flight    period 


stretches  from  April  to  September,  with  too 
small  a  sample  available  to  generalize  on  peak 
activity.  There  is  no  information  available  re- 
garding host,  collecting  techniques,  or  ecology 
of  this  species. 

DERIVATION  OF  NAME. — This  species  is 
named  in  honor  of  Dr.  L.  S.  West,  who  did  much 
of  the  early  work  with  this  genus  in  the  United 
State. 

californica  Group 
Ptilodexia  californica,  new  species 

(Figures  43-47) 

TYPE-LOCALITY. — Two  miles  (3.2  km)  S  of  Luther  Pass, 
Alpine  County,  California. 

TYPE-SPECIMENS. — The  male  holotype  and  the  allotype  are 
deposited  in  the  collection  of  the  California  Academy  of  Sci- 
ences. The  holotype  bears  the  CASC  type  number  12571. 
Complete  data  from  these  specimens  and  the  52  male  and  37 
female  paratypes  are  listed  below. 

DIAGNOSIS. — Ptilodexia  californica  adults  can 
usually  be  distinguished  from  those  of  other 
species  in  the  genus  using  the  following  combi- 
nation of  character  states:  facial  tomentum  shin- 


30 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


ing,  dark  brown  or  pale  hairs  scattered  sparsely 
along  center  of  parafacial;  haustellum  length  0.5 
to  0.65  times  head  height;  third  antennal  seg- 
ment longer  than  second,  and  length  of  plumos- 
ity  on  arista  equal  to  1.4  times  length  of  second 
antennal  segment;  frontal  vitta  obliterated  at 
narrowest  point;  scutellum  and  abdomen  of 
males  reddish,  legs  of  both  sexes  dark. 

DESCRIPTION. — Male:  Total  body  length  13.5 
mm,  dark  brown  with  abdomen  reddish  lat- 
erally. Head  height  3.3  mm;  profrons  0.3  times 
head  length.  Facial  tomentum  whitish,  shining, 
obscuring  integumental  color;  parafrontal  hairs 
long,  dark;  parafacial  hairs  dark,  rather  long, 
scattered  along  middle  of  parafacial.  Width  of 
frontal  vitta  at  antennal  base  0.5  mm,  at  narrow- 
est point,  obliterated;  carina  fairly  well  devel- 
oped, short;  facial  cavities  brown.  Height  of 
gena  0.25  times  head  height.  One  pair  of  strong 
ocellar  setae;  2  pairs  of  postocellars,  1  pair  of 
postverticals,  inner  and  outer  verticals  all  sub- 
equal.  Thirteen  pairs  of  postorbitals,  frequently 
interspersed  with  smaller  hairs;  between  post- 
orbitals and  the  whitish  occipital  hair  is  an  ir- 
regular row  of  dark  setae.  Twelve  pairs  of  fron- 
tal setae;  one  pair  of  oral  vibrissae  0.3  mm  from 
oral  margin,  the  depression  between  them  0.4 
mm  wide;  18  additional  oral  setae,  at  least  one 
above  vibrissae  and  peristomals  subequal.  Epi- 
stome  strongly  protruding,  its  width  1.6  times 
length  of  first  two  antennal  segments.  Haustel- 
lum long,  of  medium  width,  0.6  times  head 
height;  palpi  long,  narrow,  testaceous,  length  0.5 
times  haustellum  length.  Antennae  dark  rufotes- 
taceous,  third  segment,  arista  and  plumosity 
dark  brown.  Third  segment  of  antenna  1.4  times 
length  of  second  segment;  length  of  plumosity 
on  arista  1.4  times  length  of  second  antennal 
segment.  Thorax:  Mesonotum  dark  brown  with 
fine,  dull-gray  pollen,  striping  indistinct;  integ- 
ument of  scutellum  and  postalar  calli  rufotesta- 
ceous.  Six  humeral  setae,  1  posthumeral  and  2 
presuturals,  3  pairs  of  discal  scutellars.  Pleura 
dark  brown  with  fine  grayish  tomentum;  infra- 
squamal  setulae  present;  squamae  whitish, 
wings  pale  brown;  epaulet  dark  brown,  basi- 
costa  testaceous;  mediotergite  tomentose,  sub- 
shining.  Legs  dark  brown,  tibiae  rufotestaceous 
darkened  slightly  basally  and  apically;  tarsi 
blackish.  Anterior  femur  with  9  dorsal  setae,  8 
posterodorsals,  and  21  posteroventrals;  tibia 
with  dark  dorsal  line.  Middle  femur  with  3/4  an- 


terior setae  in  2  rows.  Posterior  femur  with  11 
anterodorsals,  8  anteroventrals,  5  posteroven- 
trals, and  1  anterior  seta.  Abdomen  rufous  with 
narrow,  dark-brown  median  stripe;  tomentum 
shining  white,  shining  brown  around  median 
marginals  and  posterior  median  discals  of  sec- 
ond and  third  tergites.  First  syntergite  with  1 
strong  pair  of  lateral  marginal  setae.  Second  ter- 
gite  with  3  pairs  of  median  discals,  2  pairs  of 
median  marginals,  2  pairs  of  lateral  marginals, 
and  2  pairs  of  lateral  discal  setae.  Third  tergite 
with  2  pairs  of  median  discals,  12  dorsal  and 
lateral  marginals,  and  6  pairs  of  miscellaneous 
discal  setae.  Genitalia:  Eighth  tergite  with 
sparse  long  hairs.  Ninth  tergite  with  long  brown 
hairs,  lateral  swelling  absent.  Surstylus  large 
with  lateral  angle  fairly  sharp,  depression  deep; 
surface  strongly  pitted.  Cerci  with  height  of  lat- 
eral lobes  greater  than  that  of  arms;  lateral  lobes 
slightly  attenuated  dorsally,  mesal  margins  con- 
cave; in  profile,  arms  strongly  bulging.  Ejacu- 
latory  apodeme  with  bulb  distinct  from  stem, 
which  is  twice  height  of  bulb.  Bulb  open  on  one 
side  and  on  top,  where  it  is  distinctly  lipped. 
Female:  Similar  to  male  except  for  usual  sexual 
differences  and  the  following.  Width  of  frontal 
vitta  at  narrowest  point  0.4  mm,  at  antennal  base 
0.6  mm.  Height  of  gena  0.3  times  head  height. 
Eleven  pairs  of  short  postorbital  setae;  10  pairs 
of  frontal  setae.  Legs  colored  similarly  to  those 
of  male,  tibiae  not  as  pale.  Anterior  femur  with 
14  posteroventrals,  8  posterodorsals,  and  6  dor- 
sal setae.  Middle  femur  with  5  anterior  setae  in 
2  rows.  Posterior  femur  with  no  anterior  setae, 
5  anterodorsals,  5  posteroventrals,  and  6  antero- 
ventrals. Abdomen  dark  reddish  brown;  median 
stripe  broad  and  indistinct.  First  syntergite  with 
1  pair  of  lateral  marginal  setae.  Second  tergite 
with  1  pair  of  median  marginals,  1  pair  of  median 
discals,  and  1  pair  of  lateral  marginal  setae. 
Third  tergite  with  2  pairs  of  median  discals,  2 
pairs  of  lateral  discals,  and  12  dorsal  and  lateral 
marginal  setae. 

VARIATION. — Ptilodexia  californica  adults 
show  relatively  little  variation  in  many  of  the 
diagnostic  characters  of  this  group.  However, 
they  do  seem  to  exhibit  a  distinctly  greater  than 
usual  degree  of  variation  in  minor  characters  as 
well  as  in  the  genitalia.  It  is  possible  that  later, 
more-detailed  studies  may  uncover  cryptic  or 
incipient  species  within  P.  californica.  There 
are  also  a  few  specimens  which  appear  to  be 


WILDER:  NEARCTIC  PTILODEXIA 


31 


intermediate  between  P.  californica  and  P.  sa- 
broskyi,  and  between  P.  californica  and  P.  pa- 
cifica. 

Total  body  length  ranges  from  11  to  16  mm. 
In  a  few  individuals,  facial  tomentum  has  a  yel- 
lowish cast  and  the  carina  is  keeled.  Most  spec- 
imens have  no  small  hairs  interspersed  between 
the  postorbital  setae.  The  haustellum  is  varied, 
with  the  length  ranging  from  0.5  to  0.65  times 
head  height.  Notal  tomentum  may  be  partially 
or  wholly  dull  brown  and  the  mediotergite  can 
be  fairly  shiny.  The  abdomen  is  varied  from  the 
state  shown  in  the  holotype  to  dark  reddish 
brown  with  the  median  stripe  broad  and  indis- 
tinct. The  genitalia  are  extremely  varied. 

MATERIAL  EXAMINED. — Fifty-three  males  and  39  females 
were  examined  from  the  following  localities.  California:  Al- 
pine Co.:  16,2  miles  [3.2  km]  S  Luther  Pass,  6  July  1959,  at 
flower  Achillea  Millefolium,  P.  Arnaud  (CASC);  Id,  Hope 
Valley,  9  July  1948,  J.  MacSwain  (CISC).  Fresno  Co.:  19, 
Prather,  29  June  1956,  R.  Schuster  (CISC);  299,  Watts  Val- 
ley, 22-23  June  1956,  R.  Schuster  (CISC).  Inyo  Co.:  19,  W 
Fork  Coyote  Creek,  2.1  miles  [3.4  km]  NE  Coyote  Lake, 
37°14'N,  118°30'W,  Sierra  Nevada,  9,840  ft  [ca.  3,000  m] 
sweeping,  16  Sep.  1969,  S.  Frommer  (UCRC);  Id,  near  Mono 
Pass,  12,000  ft,  [ca.  3,658  m]  19  Aug.  1956,  C.  MacNeill 
(CISC);  Id  Panamint  Mts.,  28  May  1937,  B.  Brookman 
(CASC).  Kern  Co.:  Id,  4  miles  [ca.  6.4  km]  N  Muroc,  30 
Apr.  1950,  T.  Leigh  (CISC);  2d  d,  Short  Canyon,  7  miles  [ca. 

1 1  km]  NW  Inyokern,  13  Apr.  1954,  J.  MacSwain  (CISC);  1  d , 
Walker  Pass,  11  June  1962,  G.  Bohart  (USUC).  Los  Angeles 
Co.:  3dd299,  Oct.,  Coquillett  (USNM);  5993dd,  Crystal 
Lake,  29  June  1950,  W.  Bentnick,  13d d  11 9  9,  9  July  1952, 
R.  Wagner  (CISC,  LACM);  Id,  Little  Rock,  10  May  1941,  J. 
Wilcox  (PHAC);  Idl9,  1  mile  W  Little  Rock,  13  May  1956, 
J.  MacSwain  (CISC);  1  d ,  2  miles  [ca.  3.2  km]  W  Pearblossom, 

12  May  1956,  E.  Linsley  (CISC);  19,  Rock  Creek,  21  July 
1936,  A.  Basinger  (CASC).  Mariposa  Co.:  Id,  Yosemite  Val- 
ley, 26  June  1921,  E.  Van  Dyke  (CASC).  Mono  Co.:  Idl9, 
Sardine  Creek,  27  June  1957,  D.  Flaherty  (UCRC,  UAIC); 
Id,  Sonora  Pass,  9,000-10,000  ft  [ca.  2,740-3,050  m],  16  July 
1951,  P.  Ashlock,   19,   10  Aug.   1960,  C.  Toschi  (USNM, 
CISC).  Nevada  Co. :  1  9 ,  Prosser  Dam,  15  July  1966,  D.  Miller 
(UCDC);  Id,  7  miles  [ca.  11  km]  SE  Truckee,  24  June  1954, 
G.  Schaefers  (CISC).  Plumas  Co.:  2ddl9  (Allotype),  Mead- 
ow Valley,  10  June  1924,  E.  Van  Dyke  (CASC).  Riverside 
Co.:  Id,  Hemet,  26  Apr.  1961,  Ewart  (UCRC);  Id,  Hidden 
Valley,  Joshua  Tree  National  Monument,  18  May  1946,  A. 
Melander  (USNM);  1  d ,  Pinon  Flat,  24  Apr.  1950,  C.  MacNeill 
(CISC);  Id,  Pinon  Flat,  San  Jacinto  Mts.,  18  May  1939,  E. 
Ross,  299,  Eriogonum,  21  May  1940,  H.  Reynolds,  Id,  27 
May  1939,  B.  Brookman  (CISC,  UCRC);  Id,  Pinon  Flat,  San- 
ta Rosa  Mts.,  27  May  1941 ,  D.  Knull  (OSUC).  San  Bernardino 
Co.:  Id,  Helendale,  16  May  1955,  W.  Richards  (CNCI).  San 
Diego  Co.:  19,  16.5  miles  [ca.  26.6  km]  N  of  Carlsbad,  Tim- 
berlake  (UCRC).  San  Luis  Obispo  Co.:  Id,  Oso  Flaco  Lake, 
19  July  1964,  M.  Irwin  (UCRC).  Santa  Barbara  Co.:  19,  Jan- 
ama  Beach,  9  July  1965,  J.  Powell  (CISC);  19,  Los  Prietos, 
23  June  1965,  J.  Powell  (CISC).  Santa  Cruz  Co.:  Id,  Bear 


Valley,  Santa  Cruz  Mts.,  July  1913,  F.  Clark  (CUIC).  Siskiyou 
Co.:  2d  d,  Hebron  Summit,  10  miles  [ca.  16  km]  S  of  Dorris, 
5,300  ft  lea.  1,600  m]  27  July  1953,  W.  Gertsch  (AMNH). 
Tehama  Co.:  Id399,  Deer  Creek,  6  July  1952,  M.  Cazier 
(AMNH).  Ventura  Co.:  19,  Quatal  Canyon,  NW  corner  Ven- 
tura Co.,  Stanleya  pinnata,  9  May  1959,  J.  Powell  (CISC); 
2dd,  Hungry  Valley,  5  miles  [ca.  8  km]  S  of  Gorman,  6  May 
1959,  C.  O'Brien  (CISC).  Nevada:  Clark  Co.:  19,  Kyle 
Canyon,  Charleston  Mts.,  5,200  ft  [ca.  1,580  m]  4  June  1941, 
Timberlake  (UCRC).  Ormsby  Co.:  19,  6  July,  Baker 
(USNM).  Oregon:  Hood  River  Co.:  Id,  Hood  River,  17  June, 
Childs  (CISC).  Jackson  Co.:  19,  Colestin,  31  July  1918,  E. 
Van  Duzee  (CASC).  Klamath  Co.:  Id,  Eagle  Ridge,  Klamath 
Lake,  20  May  1924,  C.  Fox  (CISC).  Washington:  Yakima  Co.: 
19,  Signal  Peak  Ranger  Stn.,  4,000  ft  [ca.  1,200  m],  15  July 
1933,  J.  Wilcox  (PHAC). 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  April  to  October  with  most  of  the  activity 
from  April  to  July.  Adults  have  been  collected 
at  low  as  well  as  high  elevations  (up  to  3,660  m). 
Flowers  from  which  they  have  been  collected 
are  Eriogonum  sp.  (Polygonaceae)  and  Stanleya 
pinnata  (Cruciferae). 

Ptilodexia  pacifica,  new  species 

(Figures  48-52) 

TYPE-LOCALITY. — Point  Pinos,  Pacific  Grove,  Monterey 
County,  California. 

TYPE-SPECIMENS.— The  holotype,  a  male,  and  the  allotype 
are  deposited  in  the  collection  of  the  California  Academy  of 
Sciences.  The  holotype  bears  the  CASC  type  number  12572. 
Complete  data  from  these  specimens  and  the  62  male  and  16 
female  paratypes  are  listed  below. 

DIAGNOSIS. — Ptilodexia  pacifica  is  a  complex 
species,  but  specimens  can  generally  be  sepa- 
rated from  those  of  other  species  in  the  genus 
by  the  following  combination  of  character 
states:  parafacial  hairs  long,  dark,  inserted  along 
center  of  parafacial  as  low  as  ventral  eye  margin; 
haustellum  length  varied,  from  0.4  to  0.55  times 
head  height;  legs  of  most  adults  either  pale  in 
color  or  marked  with  rufotestaceous;  notum  of 
most  specimens  covered  with  flat,  shiny,  brown- 
ish tomentum,  giving  it  a  coppery  appearance; 
abdominal  tomentum  concolorous — either  whit- 
ish or  brownish  in  most  individuals — white 
marked  with  brown  in  only  a  few. 

DESCRIPTION. — Male:  Total  body  length  10.5 
mm,  dark  brown,  abdomen  reddish.  Head 
height  2.75  mm;  profrons  0.25  times  head  length. 
Facial  tomentum  heavy,  shining  grayish,  ob- 
scuring integumental  color;  parafrontal  hairs 
long,  dark,  sparse;  parafacial  hairs  long,  dark, 
rather  sparse,  inserted  along  middle  of  parafacial 
to  ventral  margin  of  eye.  Width  of  frontal  vitta 


32 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


51 


FIGURES  48-52.    Ptilodexia  pacifica,  n.sp.   Fig.  48.  Genitalia  of  male,  posterior  view.   Fig.  49.  Ejaculatory  apodeme.   Fig. 
50.  Genitalia  of  male,  lateral  view.  Fig.  51.  Head  of  male,  lateral  view.  Fig.  52.  Geographical  distribution. 


at  base  of  antenna  0.4  mm,  at  narrowest  part 
obliterated;  carina  low,  broad,  rounded;  facial 
cavities  dark  brown.  Height  of  gena  0.3  times 
head  height.  One  pair  of  strong  ocellar  setae,  3 
pairs  of  postocellars,  1  pair  of  postverticals,  in- 
ner verticals,  and  outer  verticals.  Fifteen  pairs 
of  long  postorbital  setae,  not  interspersed  with 
smaller  hairs;  between  postorbitals  and  whitish 
occipital  hairs  are  a  few  irregular  dark  setae. 
Thirteen  pairs  of  frontal  setae;  1  pair  of  strong 
oral  vibrissae  0.2  mm  from  oral  margin,  the 
depression  between  them  0.4  mm  wide.  Thirteen 
additional  oral  setae;  one  above  vibrissae  and 
peristomals  subequal  to  each  other;  oral  margin 
protruding,  its  width  equal  to  2.2  times  length  of 
first  two  antennal  segments.  Haustellum  of  me- 
dium width,  length  0.5  times  head  height;  palpi 
long,  rufotestaceous,  the  length  0.6  times  haus- 
tellum  length.  Antennae  rufotestaceous,  third 
segment,  arista  and  plumosity  brown.  Third  seg- 
ment of  antenna  1 .5  times  length  of  second  seg- 
ment; length  of  plumosity  on  arista  1.75  times 
length  of  second  antennal  segment.  Thorax: 


Mesonotum  dark  brown  with  tomentum  appear- 
ing grayish  from  one  angle  and  shiny  brownish 
from  others;  striping  indistinct.  Integument  of 
postalar  calli,  scutellum,  and  sides  of  notum  ru- 
fotestaceous. Six  humeral  setae,  2  posthumerals 
and  2  presuturals,  2  pairs  of  discal  scutellars. 
Pleura  dark  brown  with  grayish  tomentum;  in- 
frasquamal  setulae  present;  squamae  white, 
wings  light  brown;  epaulet  brown,  basicosta  ru- 
fotestaceous; mediotergite  thinly  tomentose, 
shining.  Legs  dark  rufotestaceous,  anterior  and 
middle  femora  brownish  with  pale  apical  mark- 
ings; posterior  femora  with  dark  apicoventral 
patches;  tibiae  unmarked,  tarsi  dark  brown.  An- 
terior femur  with  7  dorsal  setae,  8  posterodor- 
sals,  and  15  posteroventrals;  tibia  without  dark 
dorsal  line.  Middle  femur  with  5  anterior  setae 
in  2  rows.  Posterior  femur  with  15  anterodorsals, 
8  anteroventrals,  4  posteroventrals,  and  no  an- 
terior setae.  Abdomen  rufotestaceous  with  a 
dark  median  stripe;  tomentum  whitish.  First 
syntergite  with  1  pair  of  lateral  marginal  setae. 
Second  tergite  with  2  pairs  of  median  discals,  2 


WILDER:  NEARCTIC  PTILODEXIA 


pairs  of  median  marginals,  4  pairs  of  lateral  mar- 
ginals, and  2  pairs  of  lateral  discal  setae.  Third 
tergite  with  2  pairs  of  median  discals,  10  dorsal 
and  lateral  marginals,  and  2  pairs  of  lateral  discal 
setae.  Ge  nit  alia'.  Eighth  tergite  with  sparse, 
long,  fine  hair.  Ninth  tergite  with  long  brown 
hair  and  a  distinct  lateral  swelling.  Surstylus 
with  lateral  angle  sharp,  internal  depression 
deep.  Cerci  with  lateral  lobes  longer  than  arms; 
arms  reaching  to  ventral  margin  of  surstylus;  in 
profile,  cerci  bulging,  tips  directly  ventrally. 
Ejaculatory  apodeme  with  bulb  and  stem  dis- 
tinct; stem  twice  as  long  as  height  of  bulb,  which 
is  widely  opened  on  one  side  and  top;  opposite 
side  decumbent.  Female:  Similar  to  male  except 
for  usual  sexual  differences  and  the  following: 
General  color  dark  brown.  Width  of  frontal  vitta 
at  narrowest  point  0.4  mm,  at  antennal  base  0.6 
mm  wide.  Height  of  gena  0.35  times  head  height. 
Seven  pairs  of  medium  postorbital  setae;  10 
pairs  of  frontal  setae.  Notum  dark  brown  with 
heavy  brownish  tomentum;  six  humeral  setae. 
Legs  rufotestaceous.  Anterior  femur  with  11 
posteroventrals,  4  posterodorsals,  and  5  dorsal 
setae.  Middle  femur  with  3  anterior  setae  in  2 
rows,  1  anteroventral,  and  4  posteroventrals,  4 
anteroventrals,  and  no  anterior  setae.  Abdomen 
dark  brown,  a  little  lighter  laterally.  Second  ter- 
gite with  1  pair  of  median  discals,  1  pair  of  me- 
dian marginals,  3  pairs  of  lateral  marginals,  and 
1  pair  of  lateral  discal  setae.  Third  tergite  with 
1  pair  of  median  discals,  10  dorsal  and  lateral 
marginals,  and  2  pairs  of  lateral  discal  setae.  Tip 
of  abdomen  reddish. 

VARIATION. — Ptilodexia  pacifica  is  the  most 
variable  Nearctic  speices  of  Ptilodexia.  Several 
fairly  distinct  populations  are  present,  each  of 
which  may  eventually  be  recognized  as  a  sepa- 
rate species.  I  fail  to  name  these  as  species  now 
because  of  the  inadequate  sample  and  numerous 
specimens  which  appear  to  be  intermediates. 

The  form  to  which  the  type-specimens  belong 
has  males  and  females  with  pale-colored  legs. 
These  specimens  have  the  parafacial  hairs  rather 
sparse;  are  small  in  body  size  (7-13  mm);  and 
the  abdominal  tomentum  is  uniformly  whitish  in 
most  individuals.  This  form  occurs  from  south- 
ern California  along  the  coast  into  Monterey 
County. 

Members  of  the  second  form  are  larger;  have 
parafacial  hairs  abundant;  and  only  females  have 
pale-colored  legs,  although  the  legs  of  some 
males  are  marked  with  orange  or  reddish.  The 


facial  tomentum  is  brownish  in  some  specimens, 
and  the  abdominal  tomentum  is  so  in  most. 
These  specimens  are  more  hairy  and  much  dark- 
er in  color  than  those  of  the  previous  form. 
Many  have  one  to  two  rows  of  dark  hairs  ventral 
to  the  postorbitals,  and  have  dark  squamae  and 
wings.  This  form  ranges  from  Oregon  south 
along  the  California  coast  to  Monterey  County, 
where  both  this  and  the  previous  form  are  found. 

The  third  form  is  found  in  the  Sierra  Nevada 
from  northern  California  south  into  Inyo  Coun- 
ty. These  specimens  resemble  those  of  the  first 
form  in  size  and  facial  hair;  however,  most  in- 
dividuals are  darker  in  color,  and  males  and  fe- 
males have  dark-colored  legs.  The  abdominal 
tomentum  is  concolorous,  whitish. 

Specimens  of  the  three  forms  have  a  number 
of  character  states  in  common  which  segregate 
them  from  specimens  of  other  western  species. 
It  is  for  this  reason  and  the  lack  of  material  that 
I  have  treated  the  complex  as  one  species.  It  is 
hoped  that  further  collecting,  and  ecological  and 
life  history  studies  will  elucidate  relationships 
and  uncover  a  basis  for  the  variability  within  this 
species. 

MATERIAL  EXAMINED. — Two  hundred  twenty-six  males 
and  115  females  of  P.  pacifica  were  examined.  Data  from 
those  specimens  in  the  type-series  are  as  follows:  California: 
Monterey  Co.:  35 d <J  1  9 ,  Point  Pinos,  Pacific  Grove,  24  May, 
1952,  P.  Arnaud,  4dcJ1299,  13  June  1959,  25<J<J599,  19 
Aug.  1957  (PHAC). 

Data  from  the  other  specimens  examined  are  recorded  by 
Wilder  (1976). 

DISTRIBUTION. — The  species  ranges  from 
southern  California  north  along  the  coast  to 
northern  Oregon.  It  is  also  found  in  the  Central 
Valley  and  through  the  Sierra  Nevada  in  Cali- 
fornia. 

BIOLOGICAL  NOTES. — The  flight  period  ranges 
from  May  to  November  with  specimens  occur- 
ring throughout  that  period  with  perhaps  two 
activity  peaks — one  in  late  spring  and  early  sum- 
mer, and  the  other  in  the  fall. 

Specimens  of  P.  pacifica  have  been  collected 
on  the  following  flowers  belonging  to  the  family 
Compositae:  Baccharis  pilularis,  Chrysotham- 
nus  viscidiflorus,  and  Achillea  Millefolium,  and 
undetermined  Baccharis  sp.  and  Solidago  sp. 
Specimens  have  also  been  collected  from  Eri- 
ogonum  datum  (Polygonaceae).  They  have  also 
been  collected  while  resting  on  the  leaves  of 
Rubus  parviflorus  and  Umbellularia  calif  or  nica. 
This  species  inhabits  both  low  and  high  (up  to 


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53 


FIGURES  53-57.    Ptilodexia  sabroskyi,  n.sp.  Fig.  53.  Genitalia  of  male,  posterior  view.  Fig.  54.  Ejaculatory  apodeme.  Fig. 
55.  Genitalia  of  male,  lateral  view.  Fig.  56.  Head  of  male,  lateral  view.  Fig.  57.  Geographical  distribution. 


3,660  m)  elevations.  Collecting  methods  used  in- 
clude UV  and  white  light,  sweeping,  flight  trap, 
and  light  trap.  Ptilodexia  pacifica  adults  have 
been  collected  in  meadows,  near  streams,  on 
sand  dunes,  and  on  rocks. 

Ptilodexia  sabroskyi,  new  species 

(Figures  53-57) 

TYPE-LOCALITY. — San  Clemente,  Orange  County,  Califor- 
nia. 

TYPE-SPECIMENS. — The  holotype,  a  male,  was  collected  by 
G.  Eickwort  on  25  July  1961,  and  is  deposited  in  MSUC.  The 
allotype,  deposited  in  LACM,  is  from  near  Wrightwood,  1,719 
m,  San  Bernardino  County,  California.  Complete  data  from 
these  specimens  and  the  37  male  and  7  female  paratypes  are 
listed  below. 

DIAGNOSIS. — Specimens  of  P.  sabroskyi  are 
large,  robust,  and  quite  distinctive.  They  can  be 
distinguished  from  those  of  other  species  of 
Ptilodexia  by  the  following  combination  of  char- 
acter states:  parafacial  hairs  brown,  rather 
short,  concentrated  near  upper  anterior  edge  of 
parafacial  and  scattered  randomly  elsewhere, 
lower  ones  often  yellowish;  length  of  plumosity 


on  arista  less  than  twice  length  of  second  anten- 
nal  segment,  the  third  antennal  segment  short, 
subequal  to  the  second  in  length;  smallest  dis- 
tance between  eyes  greater  than  width  of  frontal 
vitta  at  antennal  base;  frontal  vitta  not  obliter- 
ated at  its  narrowest  point;  length  of  haustellum 
varied  from  0.5  to  0.6  times  head  height;  legs  of 
both  sexes  dark;  notum  light  brown  laterally  in 
most  individuals. 

DESCRIPTION. — Male:  Total  body  length  17 
mm,  dark  brown  with  abdomen  reddish  lat- 
erally. Head  height  4  mm;  profrons  0.3  times 
head  length.  Facial  tomentum  dull  whitish, 
heavy,  obscuring  integumental  color;  parafron- 
tal  hairs  medium  in  length,  dark,  abundant; 
parafacial  hairs  brown,  short,  concentrated  on 
upper  anterior  portion  efface,  short,  pale  hairs 
scattered  elsewhere  on  parafacial.  Width  of 
frontal  vitta  at  antennal  base  0.6  mm,  at  narrow- 
est point  0.1  mm;  carina  well  developed,  broad; 
facial  cavities  dark  gray.  Height  of  gena  0.3 
times  head  height.  One  pair  of  strong  ocellar  se- 
tae, two  pairs  of  smaller  postocellars,  one  pair 


WILDER:  NEARCTIC  PTILODEXIA 


35 


each  of  postverticals,  inner  verticals,  and  outer 
verticals.  Eighteen  pairs  of  long  postorbital  se- 
tae, not  interspersed  with  smaller  hairs;  ventral 
to  postorbitals  are  no  dark  hairs,  only  yellowish 
occipital  hair.  One  pair  of  frontal  setae,  1  pair 
of  long  oral  vibrissae  0.4  mm  from  oral  margin, 
the  depression  between  them  0.6  mm  wide;  16 
additional  oral  setae,  one  above  vibrissae  rather 
strong,  peristomals  subequal  to  each  other, 
about  0.5  times  vibrissae  length.  Epistome 
strongly  protruding,  its  width  1.7  times  length  of 
first  two  antennal  segments.  Haustellum  broad, 
blackish,  length  0.5  times  head  height;  palpi 
long,  narrow,  rufotestaceous,  length  0.6  times 
haustellum  length.  Antennae  dark  rufotesta- 
ceous; third  segment,  arista,  and  plumosity  dark 
brown.  Third  segment  of  antenna  very  short,  0.8 
times  length  of  second  segment;  length  of  plu- 
mosity on  arista  1.5  times  length  of  second  an- 
tennal segment.  Thorax:  Mesonotum  dark 
brown,  rufescent  laterally;  tomentum  heavy, 
grayish,  striping  distinct;  integument  of  humeri, 
postalar  calli,  and  scutellum  rufescent.  Six  hu- 
meral setae,  2  posthumerals  and  2  presuturals, 
6  pairs  of  discal  scutellars.  Pleura  dark  brown, 
sutures  lighter,  tomentum  heavy,  grayish;  infra- 
squamal  setulae  present;  squamae  whitish, 
wings  light  brown,  epaulet  dark  brown,  basi- 
costa  testaceous;  mediotergite  subshining  to- 
mentose.  Legs  dark  brown,  tibiae  rufous,  dark- 
ened basally  and  slightly  apically;  tarsi  blackish. 
Anterior  femur  with  12  dorsal  setae,  11  postero- 
dorsals,  and  18  posteroventrals;  tibia  with  dark 
dorsal  line.  Middle  femur  with  4  strong  anterior 
setae  in  2  rows.  Posterior  femur  with  13  antero- 
dorsals,  10  anteroventrals,  6  posteroventrals, 
and  no  anterior  setae.  Abdomen  rufous  with  a 
narrow,  dark  brown,  median  stripe;  tomentum 
white,  heavy,  patchy,  brown  around  median 
marginal  setae  of  second  and  third  tergites.  First 
syntergite  with  1  pair  of  lateral  marginal  setae. 
Second  tergite  with  3  pairs  of  median  discals,  2 
pairs  of  median  marginals,  3  pairs  of  lateral  mar- 
ginals and  2  pairs  of  lateral  discals.  Third  tergite 
with  3  pairs  of  median  discals,  12  dorsal  and 
lateral  marginals,  and  4  pairs  of  lateral  discals. 
Genitalia:  Eighth  tergite  with  numerous  small, 
fine  hairs.  Ninth  tergite  with  long  dark  hairs; 
lateral  swelling  pronounced.  Surstylus  large 
with  lateral  angle  sharp,  internal  depression 
rather  shallow,  basally  strongly  convex;  in  pro- 
file, inclined  slightly  posterad.  Cerci  with  height 


of  lateral  lobes  subequal  to  height  of  arms;  in 
profile,  lateral  lobe  with  a  strong  dorsal  hook 
pointing  anteriad,  arms  bulging  slightly  ventral- 
ly,  tips  rounded.  Ejaculatory  apodeme  with  bulb 
distinct  from  long,  thin  stem,  which  is  2.5  times 
height  of  bulb.  Bulb  large  and  somewhat  flat- 
tened, open  widely  on  one  side  and  narrowly  on 
top  with  a  strong  lip;  opposite  side  strongly  de- 
cumbent. Female:  Similar  to  male  except  for 
usual  sexual  differences  and  the  following. 
Width  of  frontal  vitta  at  narrowest  point  0.4  mm, 
at  antennal  base  0.6  mm.  Height  of  gena  0.4 
times  head  height.  Nine  pairs  of  short  postor- 
bital setae  irregularly  interspersed  with  short 
hairs;  8  pairs  of  frontal  setae.  Legs  same  color 
as 'in  male,  anterior  femur  with  10  posteroven- 
trals, 8  posterodorsals,  and  8  dorsal  setae.  Mid- 
dle femur  with  3  anterior  setae  in  2  rows,  5  pos- 
teroventrals, and  1  anteroventral.  Posterior 
femur  with  no  anterior  setae,  7  anterodorsals,  4 
posteroventrals,  and  4  anteroventrals.  Abdomen 
dark  reddish  brown  with  a  vague  median  stripe; 
tomentum  heavy,  white.  First  syntergite  with  1 
pair  of  lateral  marginal  setae.  Second  tergite 
with  1  pair  of  median  discals,  1  pair  of  median 
marginals,  and  1  pair  of  lateral  marginal  setae. 
Third  tergite  with  1  pair  of  median  discals  and 
12  dorsal  and  lateral  marginals. 

VARIATION. — P.  sabroskyi  exhibits  relatively 
little  variation.  Total  body  length  ranges  from  12 
to  18  mm.  The  scattered  hairs  on  the  parafacial 
are  brown  in  some  individuals,  but  are  long  in 
none.  The  third  antennal  segment  is  varied  in 
length,  at  its  longest  being  equal  to  the  second 
segment.  The  number  of  discal  scutellar  setae  is 
quite  varied,  as  are  the  lengths  of  those  setae. 
Abdominal  color  in  males  and  females  ranges 
from  rufous  to  dark  reddish  brown,  although  few 
individuals  exhibit  the  latter.  There  is  no  brown- 
ish tomentum  on  the  abdomens  of  a  few  males 
and  many  females. 

MATERIAL  EXAMINED. — Material  examined  included  38 
males  and  8  females.  Data  from  these  specimens  are  as  fol- 
lows: California:  Unknown:  Id,  Marina  Beach,  17  June  1961, 
C.  Philip  (CASC);  16",  Pine  Lake,  southern  California,  John- 
son (USNM).  Inyo  Co.:  1$,  Wyman  Canyon,  White  Mts., 
near  stream,  21  July  1967,  S.  Frommer  (UCRC).  Los  Angeles 
Co.:  Id,  Malibu  Creek  mouth,  17  July  1953  (LACM);  Id, 
Claremont,  Baker  (LACM).  Orange  Co.:  Id,  Newport,  10 
July  1916,  D.  Hall  (USNM);  3dd,  Newport  Beach,  19  Aug. 
1920,  Timberlake  (UCRC);  Id,  San  Clemente,  25  July  1961, 
G.  Eickwort  (MSUC).  Riverside  Co.:  Id,  Coachella  Valley, 
7  Apr.  1928,  R.  Woglum  (CISC);  Id,  Palm  Springs,  Andreas 


36 


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61 


FIGURES  58-62.  Ptilodexia  carolinensis  Brauer  and  Bergenstamm.  Fig.  58.  Genitalia  of  male,  posterior  view.  Fig.  59. 
Ejaculatory  apodeme.  Fig.  60.  Genitalia  of  male,  lateral  view.  Fig.  61.  Head  of  male,  lateral  view.  Fig.  62.  Geographical 
distribution. 


Canyon,  24  Apr.  1954,  M.  Wasbauer  (CISC).  San  Bernardino 
Co.:  Id,  May,  Coquillett  (USNM);  Id,  Barton  Flats,  20  July 
1950,  A.  Melander  (USNM);  5dd,  Cajon,  11  May  1934,  C. 
M.  (UCRC);  3dd,  Colton,  26-28  May  1917,  E.  VanDuzee 
(CASC);  Id,  Deep  Creek  Public  Camp,  15  June  1957,  A. 
Menke  (LACM);  Id,  South  Fork  Camp,  San  Bernardino 
Mts.,  2  Sep.  1946,  Timberlake  (UCRC);  Id,  upper  Santa  Ana 
River,  6  July  1948,  A.  Melander,  1  d ,  9  July  1959,  1  d ,  16  July 
1947,  1  d ,  18  July  1950,  1  d  1  9 ,  22  July  1950,  1  d ,  4  Aug.  1946, 
Id,  10  Aug.  1949,  Id,  29  Aug.  1946,  Id,  15  Sep.  1946 
(USNM);  Id,  Verdemont,  22  May  1954,  A.  Melander 
(USNM);  1  9 ,  near  Wrightwood,  1,719  m  (LACM).  San  Diego 
Co.:  Id,  A.  Moldenke  (USNM);  Id,  1  mile  S  of  Del  Mar, 
Eriogonum  fasciculatum,  10  July  1963,  P.  Hurd  (CISC); 
1  d  1  $ ,  Dulzura,  14  June  1917  (AMNH);  1  9 ,  2.8  miles  [ca.  4.5 
km]  SW  of  Poway,  Los  Penasquitos  Creek,  400  ft  [ca.  120  m], 
25  July  1965,  R.  Somerby  (UCRC);  1  $,  Tecate  Peak,  10  July 
1963,  J.  Powell  (CISC).  Ventura  Co.:  19,  Oxnard  Beaches, 
13  June  1955,  R.  Erdmann,  Id,  16  July  1953,  W.  Gertsch,  19, 
25  July  1956,  R.  Erdmann  (LACM,  AMNH). 


DISTRIBUTION. — The  range  of/*,  sabroskyi  is 
restricted  to  southern  California  and  the  White 
Mountains. 

BIOLOGICAL  NOTES. — The  period  of  activity 
lasts  from  April  to  September.  Adults  have  been 


collected  near  a  stream  and  on  Eriogonum  fas- 
ciculatum (Polygonaceae). 

DERIVATION  OF  NAME. — Ptilodexia  sabroskyi 
is  named  in  honor  of  Dr.  C.  W.  Sabrosky  for  his 
enormous  contribution  to  the  study  of  the  Tach- 
inidae. 

carolinensis  Group 

Ptilodexia  carolinensis   Brauer  and  Bergen- 
stamm 

(Figures  58-62) 

Ptilodexia  carolinensis  BRAUER  AND  BERGENSTAMM,  1899:119. 
BRAUER  (1899:508);  ALDRICH  (1905:499);  SABROSKY  AND 
ARNAUD  (1965:988);  COLE  (1969:543).  [LECTOTYPE  (here 
designated),  male,  deposited  in  VMNH,  labeled,  "S.  Car- 
olina"/' 'rufipennis  Mq,  carolinensis,  Coll.  Winthem,  (two 
indistinguishable  words)' '/"Carolinensis  Type,  det.  Brauer. 
Bergenst." /"carolinensis  Type  Br  Bgst"/" Ptilodexia  caro- 
linensis Br.  Bgst."/"Lectotype  Ptilodexia  carolinensis  des- 
ignated by  D.  Wilder  1976."] 

Estheria  tibialis  (partim):  COQUILLETT  (1910:598);  TOWN- 
SEND  (1931:102).  (misidentification) 

Ptilodexia  abdominalis  (partim):  DAVIS  (1919:84).  (misidenti- 
fication) 

Ptilodexia  neotibialis  WEST,  1924:184.  LEONARD  (1928:822); 


WILDER:  NEARCTIC  PTILODEXIA 


37 


CURRAN  (1930:93);  SABROSKY  AND  ARNAUD  (1%5:989); 
COLE  (1969:543).  [HOLOTYPE,  male,  deposited  in  MCZC, 
labeled,  "Colebrook,  Conn.,  Coll.  W.  M.  Wheeler'VBlack 
square/"MCZ  Type  26963 "/"Holotype  Ptilodexia  neotibi- 
alis  West."]  NEW  SYNONYMY 

Dexia  harpasa  (partim):  ALDRICH  (1925:114);  JOHNSON 
(1925b:208).  (misidentification) 

Rhynchodexia  confusa  (partim):  JOHNSON  (1925b:208).  (mis- 
identification) 

Ptilodexia  minor  WEST,  1925: 132.  LEONARD  (1928:822);  WEST 
(1950:pl.  I,  fig.  3,  pi.  IV,  fig.  8);  SABROSKY  AND  ARNAUD 
(1965:989).  [HOLOTYPE,  female,  deposited  in  CUIC,  labeled 
"Duck  Lake,  N.Y.,  6-VHI-21"/"L.  S.  West  Collector"/ 
"Wing  Slide,  Cornell  U.,  Lot  919,  Sub  86,  L.  S.  West"/ 
"Holotype  Ptilodexia  minor  West"/"Holotype  Cornell  U., 
No.  1875'V 'Cornell  U.,  Lot.  922,  Sub.  43."]  NEW  SYN- 
ONYMY 

TAXONOMIC  NOTES. — Ptilodexia  carolinensis 
has  been  confused  routinely  with  other  species 
by  earlier  workers.  Neither  Curran  nor  West  had 
ever  seen  the  type.  West  speaks  of  generic  char- 
acters but  never  mentions  P.  carolinensis,  the 
type-species  of  the  genus,  in  any  of  his  papers. 

Aldrich  (1925)  saw  the  types,  matched  them 
with  his  specimens  of  P.  canescens  and  P.  plan- 
ifrons,  which  he  thought  were  Estheria  tibialis. 
Following  Austen  (1907),  he  assigned  this 
species  to  Dexia  harpasa  Walker.  Townsend 
(1931)  and  Coquillett  (1910)  also  felt  that  P.  car- 
olinensis was  a  synonym  of  £.  tibialis.  Sabrosky 
and  Arnaud  (1965)  considered  P.  carolinensis 
separate  from  P.  harpasa. 

I  have  seen  the  type — there  is  only  one  male 
now,  not  the  two  which  Aldrich  (1925)  report- 
ed— and  it  is  in  excellent  condition.  The  speci- 
men has  been  designated  lectotype. 

DIAGNOSIS. — Ptilodexia  carolinensis  is  a  di- 
verse species  (or  perhaps  species  group),  its 
members  lacking  distinct  characters.  Specimens 
can  be  more  easily  assigned  here  by  character- 
istics which  they  lack  rather  than  those  which 
they  possess. 

Adults  with  rather  broad,  parallel-sided  face, 
vibrissal  angle  protruding  little  or  not  at  all; 
parafacial  hairs  long,  fine,  inserted  along  middle 
of  parafacial;  epistome  protruding  only  slightly; 
third  antennal  segment  1.5  times  length  of  sec- 
ond in  most  specimens;  length  of  plumosity  on 
arista  varied  from  1  to  1.5  times  length  of  second 
antennal  segment;  haustellum  length  varied  from 
0.4  to  0.5  times  head  height;  scutellum  and  sides 
of  abdomen  orange-brown  in  most  adults;  some 
specimens  from  Texas  and  South  Dakota  are  to- 
tally orange-brown. 


MATERIAL  EXAMINED. — Two  hundred  fifty- 
six  males  and  133  females  were  examined. 

DISTRIBUTION. — This  species  ranges  widely 
from  Nova  Scotia  south  to  Georgia  and  west  in- 
to Texas  and  the  Rocky  Mountains,  with  scat- 
tered records  from  Utah,  Wyoming,  and  Sas- 
katchewan. 

BIOLOGICAL  NOTES. — The  flight  period  is 
from  July  to  September  with  the  majority  of 
specimens  collected  from  mid-August  to  mid- 
September.  Specimens  have  been  collected  as 
early  as  30  May  on  Long  Island  and  as  late  as 
23  October  in  Virginia. 

In  Texas,  the  insect  seems  to  be  double- 
brooded,  some  specimens  being  collected  in 
April,  May,  and  June,  and  others  in  September, 
October,  and  November.  In  material  from  other 
areas,  such  as  New  York,  we  find  early-season 
and  late-season  specimens,  but  their  numbers  do 
not  indicate  a  double  brood.  There  may  be  a 
correlation  between  the  brood  and  body  color 
in  the  enigmatic  Texas  specimens.  Unfortu- 
nately, the  sample  at  hand,  while  giving  weight 
to  this  possibility,  is  too  small  for  any  such  con- 
clusions to  be  drawn. 

Specimens  which  have  been  brought  into  the 
lab  complete  their  development  and  emerge 
much  earlier  than  they  would  have  in  nature. 
Parasitized  Japanese  beetles,  collected  in  Oc- 
tober, yielded  adult  Ptilodexia  carolinensis  as 
early  as  21  December.  It  is  not  known  if  these 
specimens  were  exposed  to  cold  temperatures. 
It  is  feasible  that  in  a  warm  climate  two  broods 
might  occur.  In  the  area  where  the  above-men- 
tioned specimens  were  collected  (Connecticut), 
the  adult  would  not  have  emerged,  under  normal 
conditions,  until  the  following  summer. 

Nearly  all  of  the  specimens  of  this  species 
were  collected  at  low  elevations.  One  was  col- 
lected at  1,450  m  in  the  southern  Appalachians, 
and  others  in  Colorado  and  Wyoming  at  1,950 
m  and  2,060  m,  respectively.  A  number  of  spec- 
imens have  been  collected  by  sweeping  vegeta- 
tion, and  some  have  been  taken  on  Solidago  sp. 
Perhaps  coincidentally,  Japanese  beetle  feeds  on 
the  foliage  of  Solidago  juncea  (Fleming  1972). 
Available  host  data  indicate  that  this  insect  par- 
asitizes Popillia  japonica  in  the  Northeast  and 
Phyllophaga  sp.  in  Texas.  Davis  (1919:84)  re- 
ports that  adults  (under  the  name  of  P.  abdom- 
inalis  were  reared  from  the  larvae  of  Phyllopha- 
ga rugosa  collected  in  Manitoba. 


38 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


FIGURES  63-67.    Ptilodexia  canescens  (Walker).    Fig.  63.  Genitalia  of  male,  posterior  view.    Fig.  64.  Ejaculatory  apo- 
deme.  Fig.  65.  Genitalia  of  male,  lateral  view.  Fig.  66.  Head  of  male,  lateral  view.  Fig.  67.  Geographical  distribution. 


Ptilodexia  canescens  (Walker) 

(Figures  63-67) 

Dexia  canescens  WALKER,  1852:310.  OSTEN  SACKEN 
(1878:155);  ALDRICH  (1905:502);  AUSTEN  (1907:345); 
CHAMPLAIN  AND  KNULL  (1944:214);  PETERSON  (1948:60); 
SABROSKY  AND  ARNAUD  (1965:988).  [HOLOTYPE,  male,  de- 
posited in  BMNH,  labeled  "Dexia  canescens,  Walk."/ 
••Holotype'VU.S. "/"United  States.  Ex.  Coll.  Saunders 
68.4"/Crosskey  examination  label,  1970] 

Ptilodexia  harpasa  (partim):  DAVIS  (1919:82);  ALDRICH 
(1925:114);  JOHNSON  (1925b:208).  (misidentification) 

Rhynchodexia  confusa  (partim):  JOHNSON  (1925b:208).  (mis- 
identification) 

TAXONOMIC  NOTES. — The  holotype  has  many 
dorsal  setae  broken  and  one  antenna  missing. 
The  wings  are  in  excellent  condition. 

DIAGNOSIS. — Adults  of  P.  canescens  are 
unique  in  the  following  combination  of  charac- 
ters: parafacial  hairs  long,  coarse,  denser  near 
lower  anterior  edge  of  eye  in  most  individuals. 
Oral  margin  in  most  adults  not  protruding;  males 
with  frontal  stripe  not  obliterated  at  narrowest 
point;  width  of  plumosity  on  arista  less  than  or 
equal  to  length  of  second  antennal  segment; 
length  of  haustellum  less  than  half  head  height; 


total  absence  of  infrasquamal  setulae;  abdomen 
quite  dark  in  most  specimens,  with  a  distinct 
median  stripe  in  a  few. 

MATERIAL  EXAMINED. — One  hundred  forty- 
eight  males  and  68  females  were  examined. 

DISTRIBUTION. — This  species  ranges  from 
Nova  Scotia  to  Long  Island  and  west  to  the 
Rocky  Mountains.  There  are  scattered  records 
in  British  Columbia,  Idaho,  Wyoming,  and 
Utah. 

BIOLOGICAL  NOTES. — The  flight  period  is 
from  June  to  August,  with  the  majority  of  rec- 
ords in  mid-July.  Specimens  have  been  collected 
as  early  as  24  April  (Long  Island,  1923)  and  as 
late  as  8  September  (Massachusetts,  1907)  in 
nature. 

The  majority  of  specimens  were  collected  at 
low  elevations;  however,  there  are  records  from 
mountainous  areas,  the  highest  being  2,040  m  in 
Wyoming.  Specimens  from  Massachusetts  and 
Utah  have  been  collected  on  the  flowers  of  par- 
snip. Specimens  from  State  College,  Pennsyl- 
vania, were  taken  at  a  UV  light.  In  Kindred,  in 
eastern  North  Dakota,  a  male  and  a  female  were 


WILDER:  NEARCTIC  PTILODEXIA 


FIGURES  68-72.    Ptilodexia  halone  (Walker).  Fig.  68.  Genitalia  of  male,  posterior  view.  Fig.  69.  Ejaculatory  apodeme.  Fig. 
70.  Genitalia  of  male,  lateral  view.  Fig.  71.  Head  of  male,  lateral  view.  Fig.  72.  Geographical  distribution. 


collected  on  native  prairie  vegetation.  Two 
males  and  six  females  have  been  reared  and  are 
associated  with  puparia.  The  only  host  data 
available  indicate  that  this  insect  is  a  parasitoid 
of  Phyllophaga  sp. 

Ptilodexia  halone  (Walker) 

(Figures  68-72) 

Dexia  halone  WALKER,  1849:837.  OSTEN  SACKEN  (1878:155); 
ALDRICH  (1905:502);  SABROSKY  AND  ARNAUD  (1965:988). 
[LECTOTYPE  (here  designated),  male,  desposited  in  BMNH, 
labeled,  "Dexia  halone  Wlk."/"Lectotype"/"Georgia, 
U.S.A.,  Ex.  Coll.  Abbot"/" Georgia'VCrosskey  examina- 
tion label  1970/"Lectotype  Dexia  halone  Walker  designated 
by  D.  Wilder  1975. "J 

Ptilodexia  hucketti  WEST,  1952:131.  LEONARD  (1928:822); 
CURRAN  (1930:93);  WEST  (1950:pl.  I,  fig.  1);  SABROSKY  AND 
ARNAUD  (1965:988)  [HOLOTYPE,  male,  deposited  in  CUIC, 
labeled,  "Riverhead,  L.I.,  N.Y.,  30  July  1922,  H.  C.  Huck- 
ett'V'Holotype  Ptilodexia  hucketti  West,  d'VHolotype 
Cornell  U.  No.  1872'VCornell  U.,  Lot.  922,  Sub.  44"| 
NEW  SYNONYMY 

TAXONOMIC  NOTES. — The  lectotype  and 
paralectotype  are  in  poor  condition  but  still  eas- 
ily recognizable.  The  lectotype  has  all  dorsal  se- 
tae broken  or  absent,  most  tarsi  and  the  left  pos- 


terior leg  missing,  and  the  abdomen  broken  and 
glued. 

DIAGNOSIS. — Adults  of  Ptilodexia  halone  can 
be  distinguished  from  those  of  all  other  species 
of  Ptilodexia  by  the  following  combination  of 
characters:  totally  orange  or  orange-brown  in 
color;  parafacial  hairs  short  and  pale,  inserted 
rather  randomly  on  parafacial;  oral  margin  not 
or  only  slightly  protruding;  area  between  oral 
vibrissae  and  oral  margin  higher  than  wide 
(length  and  width  may  be  equal);  length  of  haus- 
tellum  0.3  to  0.4  times  head  height;  third  anten- 
nal  segment  only  slightly  longer  than  second; 
length  of  plumosity  on  arista  1.4  times  length  of 
second  segment;  inner  vertical  setae  strongly 
convergent;  tarsi  concolorous  with  femora  and 
tibiae;  presence  of  median  marginal  setae  on 
first  abdominal  segment. 

MATERIAL  EXAMINED. — Thirteen  males  and 
seven  females  were  examined. 

DISTRIBUTION. — Scant  available  records  show 
that  P.  halone  ranges  from  southern  Mississippi 
presumably  through  the  Appalachians  into  Ten- 
nessee and  along  the  East  Coast  from  Maryland 


40 


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76 


FIGURES  73-77.    Ptilodexia  prexaspes  (Walker).   Fig.  73.  Genitalia  of  male,  posterior  view.  Fig.  74.  Ejaculatory  apodeme. 
Fig.  75.  Genitalia  of  male,  lateral  view.  Fig.  76.  Head  of  male,  lateral  view.  Fig.  77.  Geographical  distribution. 


to  Albany,  New  York.  There  is  one  record  from 
Georgia.  Adults  appear  to  be  uncommon. 

BIOLOGICAL  NOTES. — The  flight  period  ranges 
from  June  to  July  in  the  southern  part  of  its 
range  and  from  mid-July  to  late  August  in  the 
north.  There  are  no  ecological  data  associated 
with  the  material  examined. 

Ptilodexia  prexaspes  (Walker) 

(Figures  73-77) 

Dexia  prexaspes  WALKER,  1849:837.  OSTEN  SACKEN  (1878:155); 
BRAUER  (1899:502);  AUSTEN  (1907:345);  BRIMLEY  (1938:368); 
SABROSKY  AND  ARNAUD  (1965:989).  [LECTOTYPE  (here  des- 
ignated), male,  deposited  in  BMNH,  labeled,  "Dexia  prex- 
aspes Walk."/"Lectotype"/"Georgia,  U.S.A.,  Ex.  coll. 
Abbot"/"Georgia"/Crosskey  examination  label,  1970/ 
"Lectotype  Dexia  prexaspes  Walker,  designated  by  D. 
Wilder  1975."] 

TAXONOMIC  NOTES. — The  lectotype  has  the 
right  front  and  rear  tarsi  missing,  most  frontals, 
dorsal  abdominal,  and  thoracic  setae  missing, 
and  the  haustellum  withdrawn. 

DIAGNOSIS. — Adults  of  Ptilodexia  prexaspes 
are  unique  in  the  following  combination  of  char- 
acters: Rather  light-colored  flies,  females  uni- 


formly dark  rufotestaceous  except  for  darker 
tarsi;  males  may  be  partially  brown  dorsally,  but 
with  at  least  humeri  and  legs  pale  rufotesta- 
ceous; parafacial  hairs  dark,  short  in  females, 
long  and  coarse  in  males,  scattered  on  parafa- 
cial; third  antennal  segment  1.5  times  length  of 
second;  length  of  plumosity  on  arista  subequal 
to  second  antennal  article;  frontal  vitta  of  males 
obliterated  at  some  point  between  eyes;  length 
of  haustellum  0.3  to  0.4  times  head  height;  oral 
margin  protrudes  slightly,  if  at  all;  area  between 
oral  vibrissae  and  oral  margin  distinctly  higher 
than  wide;  legs  of  males  and  females  light  in 
color. 

MATERIAL  EXAMINED. — Only  six  males  and 
six  females  of  P.  prexaspes  have  been  examined 
for  this  study. 

DISTRIBUTION. — This  species  appears  to  be 
localized  and  its  members  uncommon.  The 
range  extends  from  central  Florida  along  the 
southern  Atlantic  coast  to  Cape  Hatteras,  North 
Carolina. 

BIOLOGICAL  NOTES. — This  insect  appears  to 
have  two  broods  in  Florida.  The  flight  periods 


WILDER:  NEARCTIC  PTILODEXIA 


41 


FIGURES  78-82.    Ptilodexia  harpasa  (Walker).   Fig.  78.  Genitalia  of  male,  posterior  view.   Fig.  79.  Ejaculatory  apodeme. 
Fig.  80.  Genitalia  of  male,  lateral  view.  Fig.  81.  Head  of  male,  lateral  view.  Fig.  82.  Geographical  distribution. 


are  from  late  March  and  April  to  June  in  Florida, 
and  late  August  and  September  to  October  in 
Florida  and  North  Carolina. 

Adults  of  P.  prexaspes  have  been  collected  at 
a  15-watt  UV  light  in  March  and  June.  In  the 
fall  they  have  been  taken  on  Solidago  micro- 
cephala  and  Eupatorium,  both  Compositae. 
One  specimen  was  collected  from  a  female  asi- 
lid,  Efferia  aestuans  (Linnaeus),  which  had  pre- 
sumably taken  it  as  prey. 

harpasa  Group 
Ptilodexia  harpasa  (Walker) 

(Figures  78-82) 

Dexia  harpasa  WALKER,  1849:840.  OSTEN  SACKEN  (1878:155); 
ALDRICH  (1905:502);  AUSTEN  (1907:344);  DAVIS  (1919:82); 
WEST  (1924: 186);  ALDRICH  (1925: 1 14);  JOHNSON  (1925a:89); 
(1925b:208);  (1927:203);  LEONARD  (1928:822);  CURRAN 
(1930:93);  TOWNSEND  (1931:102);  BRIMLEY  (1938:368); 
PROCTER  (1938:371);  (1946:429);  SABROSKY  AND  ARNAUD 
(1965:988).  [HOLOTYPE,  a  female,  deposited  in  BMNH,  la- 
beled, "Dexia  harpasa,  Walk."/"Holotype"/"40  3  30 
1026"/"N.  America,  ex.  Coll.  Children  40.3.30. 1026."/ 
Crosskey  examination  label  1970.] 

Ptilodexia  leucoptera  WEST,  1925:132.  LEONARD  (1928:822); 
CURRAN  (1930:93);  WEST  (1950:pl.  I,  fig.  2);  SABROSKY  AND 


ARNAUD  (1965:989).  [HOLOTYPE,  female,  deposited  in 
CUIC,  labeled,  "Riverhead,  L.I.,  N.Y.,  IX-17-13"/"Wing 
Slide,  Cornell  U.,  Lot.  919,  Sub.  115,  L.  S.  West'VHo- 
lotype  Ptilodexia  leucoptera  West  9"/"Holotype  Cornell 
U.,  No.  1874"/"Cornell  U.,  Lot.  922,  Sub.  42."]  NEW  SYN- 
ONYMY 

Dinera  robusta  CURRAN,  1930:93.  WEST  (1950:111);  SABROS- 
KY AND  ARNAUD  (1965:989).  [LECTOTYPE  (here  designat- 
ed), male,  deposited  in  CUIC,  labeled,  "Poughkeepsie, 
N.Y.July  1906,  N.Y.S.  Coll."/" d  Allotype  Rhynchiodexia 
robusta  Curran'VAllotype  Cornell  U.,  No.  1935'VCor- 
nell  U.,  Lot.  922,  Sub.  V' I" Ptilodexia  obscura  West,  det. 
H.  J.  Reinhard'VLectotype  Dinera  robusta  Curran  des- 
ignated by  D.  Wilder  1976."]  NEW  SYNONYMY 

Ptilodexia  tibialis  (partim):  TOWNSEND  (1931:102). 

TAXONOMIC  NOTES. — Ptilodexia  harpasa  was 
first  described  and  named  from  one  female  by 
Walker  in  1849.  The  holotype  is  in  poor  condi- 
tion, lacking  posterior  legs,  right  middle  leg,  and 
antennae  (one  antenna  is  glued  to  the  double 
mount).  All  the  setae  from  the  dorsal  surface  of 
the  thorax  and  abdomen  are  missing,  and  the 
coxal,  sternopleural,  and  hypopleural  areas  of 
the  left  side  are  obliterated  by  the  pin.  The  para- 
facial  hairs  have  been  rubbed  off,  but  were  ob- 
viously present  at  one  time.  Despite  its  condi- 


42 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


tion,  there  is  no  question  about  the  identity  of 
the  specimen. 

Austen  (1907)  placed  the  name  P.  harpasa  in 
synonymy  with  P.  tibialis,  apparently  without 
seeing  the  type  of  the  latter  (which  had  been 
lost).  Aldrich  (in  Davis  1919)  disputed  this  and 
restored  validity  to  P.  harpasa.  In  1925,  upon 
visiting  the  Vienna  Natural  History  Museum,  he 
placed  P.  carolinensis  in  synonymy  with  P.  har- 
pasa, using  for  comparison  specimens  from 
USNM  which  were  actually  P.  canescens  and 
P.  planifrons.  At  no  time  does  Aldrich  mention 
seeing  the  type  of  P.  harpasa;  he  apparently 
based  his  conclusions  solely  on  the  word  of  Aus- 
ten. In  1931,  Townsend  reiterated  this  synony- 
my and  restored  the  name  P.  tibialis  as  the  valid 
one,  again  without  looking  at  the  latter  type. 
Since  that  time,  the  names  P.  tibialis  and  P. 
harpasa  have  both  been  used  for  many  species 
of  Ptilodexia. 

Ptilodexia  leucoptera  was  also  described  from 
a  single  female  by  West  in  1925.  The  holotype, 
from  Riverhead,  Long  Island,  is  in  good  condi- 
tion and  is  deposited  in  CUIC. 

Curran  introduced  the  name  Dinera  robusta 
into  the  literature  in  his  1930  key.  West  (1950) 
claimed  that  he  had  never  intended  to  describe 
robusta  in  the  genus  Dinera.  This  manuscript 
name  was  only  tentative — until  Curran  validated 
it.  The  specific  epithet,  robusta,  is  totally  mis- 
leading since  this  species  is  one  of  the  smallest 
in  the  genus.  West  redescribed  and  illustrated 
Rhynchiodexia  robusta  in  1950,  incorrectly  des- 
ignating his  earlier  types  as  holotype  and  allo- 
type.  Since  Curran  was  the  describer,  the  spec- 
imens he  looked  at  (including  West's  holotype 
and  allotype)  became  a  syntype  series,  and  the 
only  valid  designation  could  be  lectotype  and 
paralectotype.  I  have  designated  West's  female 
"holotype"  as  paralectotype  and  his  male  "al- 
lotype" as  lectotype.  Other  determined  material 
in  AMNH  and  CUIC  which  Curran  probably 
used  in  developing  his  concept  of  the  species 
have  also  been  designated  paralectotypes. 

DIAGNOSIS. — Ptilodexia  harpasa  adults  are 
among  the  smallest,  most  slender  flies  in  the  ge- 
nus. They  have  parafacial  hairs  medium  to  long, 
fine,  distributed  sparsely  along  center  of  para- 
facial,  in  a  few  individuals  inserted  below  ven- 
tral eye  margin;  haustellum  length  0.65  to  0.7 
times  head  height,  slender;  third  antennal  seg- 
ment long;  length  of  plumosity  on  arista  more 


than  twice  length  of  second  antennal  segment; 
abdomen  and  scutellum  appear  dark,  but  are  ac- 
tually partially  rufescent. 

MATERIAL  EXAMINED. — Four  hundred  seven 
males  and  465  females  were  examined,  a  sex 
ratio  different  from  the  usual  in  the  genus. 

DISTRIBUTION. — Ptilodexia  harpasa  ranges 
from  the  southern  Appalachians  into  northern 
Quebec  and  west  into  Ohio,  Michigan,  northern 
Illinois,  Minnesota,  North  Dakota,  and  Mani- 
toba. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  May  to  October,  the  most  active  months 
being  August  and  September.  Ptilodexia  har- 
pasa adults  have  been  collected  in  Malaise 
traps,  by  net,  on  foliage,  and  as  the  prey  of  a 
crab  spider.  Flowers  from  which  they  have  been 
collected  are:  Solidago  canadensis,  Solidago 
rugosa,  Eupatorium  coelestinum  (Compositae); 
and  Melilotus  alba  (Fabaceae). 

Several  specimens  have  been  reared  and  are 
pinned  with  puparia.  Two  specimens  from  On- 
tario bear  the  label,  "reared  from  R.  C.  larvae." 
This  is  perhaps  an  abbreviation  for  rose  chafer, 
Macrodactylus  subspinosus,  a  small  scarab, 
whose  range  coincides  with  that  of  P.  harpasa. 
Davis  (1919)  reports  that  adults  have  been 
reared  from  grubs  (Phyllophaga?)  from  Con- 
necticut. 

Ptilodexia  rufipennis  (Macquart) 

(Figures  83-88) 

Dexia  rufipennis  MACQUART,  1842:244.  OSTEN  SACKEN 
(1878:155);  BRAUER  (1899:508);  JOHNSON  (1925b:209);  AL- 
DRICH (1905:504);  CURRAN  (1930:93);  BRIMLEY  (1938:367); 
SABROSKY  AND  ARNAUD  (1965:989).  [HOLOTYPE,  female, 
lost.] 

Dexia  cerata  WALKER,  1849:847.  OSTEN  SACKEN  (1878:155); 
ALDRICH  (1905:502);  AUSTEN  (1907:344);  FATTIG  (1944:3); 
SABROSKY  AND  ARNAUD  (1965:989).  [HOLOTYPE,  female, 
deposited  in  BMNH,  labeled,  "Dexia  cerata  Walk.'T'Ho- 
lotype'V'N.  Amer.'V'N.  America.  Pres.  by  the  Entomo- 
logical Club,  44.12"/Crosskey  examination  label,  1970.] 
NEW  SYNONYMY 

Dexia  albifrons  WALKER,  1852:317.  OSTEN  SACKEN  (1878:155); 
ALDRICH  (1905:501);  AUSTEN  (1907:345);  SABROSKY  AND 
ARNAUD  (1965:988).  [HOLOTYPE,  male,  deposited  in 
BMNH,  labeled,  "Dexia  albifrons  Walk. "/"Holotype"/ 
"U.S. "/"United  States  Ex.  Coll.  Saunders.  68.4"/"a/W- 
/raws  "/Crosskey  examination  label,  1970.]  NEW  SYNONY- 
MY 

Rhynchodexia  confusa  WEST,  1924:185.  JOHNSON  (1925b:208); 
LEONARD  (1928:821);  CURRAN  (1930:93);  WRAY  (1938:91); 
REINHARD  (1943:22);  WEST  (1950:109);  SABROSKY  AND  AR- 
NAUD (1965:988).  [HOLOTYPE,  male,  deposited  in  CUIC, 
labeled,  "Selden,  L.I.,  July  1,  1923,  N.Y. "/"Holotype 


WILDER:  NEARCTIC  PTILODEX1A 


43 


83 


FIGURES  83-88.  Ptilodexia  rufipennis  (Macquart).  Fig.  83.  Genitalia  of  male,  posterior  view.  Fig.  84.  Ejaculatory  apo- 
deme.  Fig.  85.  Ejaculatory  apodeme.  Fig.  86.  Genitalia  of  male,  lateral  view.  Fig.  87.  Head  of  male,  lateral  view.  Fig.  88. 
Geographical  distribution. 


Rhynchodexia  confusa  West,  d"/"Holotype,  Cornell  U. 
No.  1245'V 'Cornell  U.,  Lot.  922,  Sub.  35."]  NEW  SYN- 
ONYMY 

Rhynchodexia  translucipennis  WEST,  1925:135.  LEONARD 
(1928:821);  CURRAN  (1930:93);  WEST  (1950:pl.  I,  fig.  6);  SA- 
BROSKY  AND  ARNAUD  (1965:989).  [HOLOTYPE,  male,  de- 
posited in  CUIC,  labeled,  "Ithaca,  N.Y.,  18  July  'Zl'VL. 
S.  West  Collector"/1 'Wing  Slide,  Cornell  U.,  Lot  919,  Sub. 
116,  L.  S.  West'Y'Holotype,  Rhynchodexia  translucipen- 
nis West,  cTVHolotype  Cornell  U.,  No.  826"/"Cornell 
U.,  Lot  922,  Sub.  37. "J  NEW  SYNONYMY 

Ptilodexia  harpasa  (partim):  JOHNSON  (1925b:209).  (misiden- 
tification) 

Ptilodexia  neotibialis  (partim):  JOHNSON  (1925:208).  (misiden- 
tification) 

Rhynchiodexia  dubia  CURRAN,  1930:93.  WEST  (1950: 110);  SA- 
BROSKY  AND  ARNAUD  (1965:988).  [LECTOTYPE  (here  des- 
ignated), male,  deposited  in  AMNH,  labeled,  "Sta.  Study 
Insects,  Tuxedo,  N.Y.,  28-VII-1928"/"Collector  C.  H. 
Curran'VLectotype  Rhynchiodexia  dubia  Curran  desig- 
nated by  D.  Wilder  1976."]  NEW  SYNONYMY 

TAXONOMIC  NOTES. — The  name  P.  rufipennis 
has  been  misused  frequently  in  the  literature.  It 
also  has  a  long  list  of  synonyms.  Part  of  the 
reason  for  this  is  clear — the  species  exhibits  ex- 
ceptionally high  variability.  Females  and  males 


differ  from  each  other  in  taxonomic  characters 
which  were  used  previously  in  the  group  to  de- 
fine species.  There  is  also  strong  clinal  variation 
in  males,  which  without  being  thoroughly  sam- 
pled could  lead  workers  to  name  certain  popu- 
lations as  separate  species.  Specimens  also  are 
commonly  collected.  These  factors,  combined 
with  the  failure  of  earlier  workers  to  examine 
type  material,  have  contributed  to  the  taxonom- 
ic problems  in  this  species. 

The  name  has  been  in  general  use  since  it  was 
proposed  in  1843.  Brauer  (1899)  felt  that  this 
species  was  the  same  as  P.  carolinensis,  and 
therefore  placed  the  species  in  Ptilodexia.  Sub- 
sequent workers  used  the  name  but  failed  to  de- 
fine the  species.  This  was  understandable  since 
the  type  was  lost  and  the  original  description 
was  short. 

Curran's  1930  key  characterized  members  of 
the  species  as  having  infrasquamal  setulae  ab- 
sent, parafacials  haired,  and  the  scutellum  red- 
dish. Macquart's  description  states  that  the  fe- 
male from  Nova  Scotia,  which  he  named  Dexia 


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rufipennis,  has,  among  other  characteristics,  the 
legs,  basal  antennal  segments,  and  scutellum 
"testace."  He  also  describes  the  epistome  as 
"saillant"  or  protruding.  The  only  species  which 
ranges  into  Nova  Scotia  and  whose  members 
combine  those  characters  with  the  others  in 
Macquart's  original  description  is  the  one  pres- 
ently being  considered,  specimens  of  which  lack 
parafacial  hairs. 

The  holotype  of  Dexia  cerata  Walker,  de- 
scribed as  being  from  "North  America,"  fits 
Macquart's  description  perfectly.  It  is  in  poor 
condition,  with  antennae,  facial  setae,  and  left 
middle  and  hind  legs  broken.  The  haustellum  is 
withdrawn. 

Dexia  albifrons  Walker,  another  synonym  of 
this  species,  is  also  from  the  "United  States." 
The  male  holotype  is  a  dark,  small  specimen  of 
this  varied  species.  It  is  in  poor  condition,  en- 
tirely covered  with  dust  or  mold  and  missing 
most  setae. 

West  used  the  genus  Rhynchodexia  for  those 
species  whose  members  lacked  parafacial  hairs. 
In  describing  Rhynchodexia  confusa,  he  looked 
only  at  specimens  from  the  northeastern  United 
States  and  suspected  that  more  than  one  species 
was  involved.  He  also  stated  that  this  species 
had  been  represented  in  collections  under  the 
name  P.  harpasa,  another  Walker  species  de- 
scribed in  the  same  paper  as  P.  cerata.  The  de- 
scription of  P.  harpasa  is  much  less  character- 
istic of  members  of  this  species  than  is  that  of 
P.  cerata,  and  it  is  difficult  to  understand  why 
that  name  was  being  used  instead  of  P.  cerata. 
The  holotype  and  allotype  of  P.  confusa  are 
both  large  specimens  with  light-colored  legs. 
They  are  in  excellent  condition. 

West's  other  synonym  of  P.  rufipennis  is 
Rhynchodexia  translucipennis.  This  species  was 
described  from  two  unusual-looking  specimens 
of  P.  rufipennis.  The  holotype  (and  paratype) 
has  translucent,  milky- white  wings  and  a  light, 
reddish-brown  thorax.  It  is  possible  that  these 
specimens  are  teneral;  however,  they  lack  the 
characteristic  collapsed  face  normally  associ- 
ated with  teneral  Ptilodexia.  Tenerals  do  not 
generally  have  milky  wings.  Nevertheless,  these 
specimens  definitely  belong  in  P.  rufipennis. 
The  type-specimens  are  in  good  condition. 

The  name  Rhynchiodexia  dubia  was  intro- 
duced into  the  literature  by  C.  H.  Curran  in 
1931.  Between  1921  and  1925,  West  had  been 
working  with  tachinids  of  New  York  and  adja- 


cent states  and  had  placed  type  labels  on  certain 
specimens.  His  work  was  interrupted,  and  these 
specimens,  with  their  manuscript  names,  were 
deposited  in  CUIC.  Curran  consulted  this  col- 
lection and  used  five  of  West's  manuscript 
names  in  his  paper  "Diptera  from  Tuxedo," 
which  was  essentially  a  faunal  list.  Unfortu- 
nately, a  key  to  adults  of  the  species  of  Rhyn- 
chiodexia appears  in  this  paper,  making  the 
names  valid  since  they  were  accompanied  by  a 
statement  of  diagnostic  characters. 

Of  the  five  species  described  in  this  paper, 
Curran  mentions  actual  material  for  only  one — 
P.  dubia.  He  does  not  mention  West's  holotype 
(a  female  from  Cimarron,  Colorado),  although 
he  must  have  seen  it  to  use  the  name.  West's 
holotype  belongs  to  P.  agilis  Reinhard;  the 
specimens  which  Curran  mentions  belong  to  P. 
rufipennis  and  are  from  New  York.  I  have  des- 
ignated the  lectotype  of  P.  dubia  from  the  Cur- 
ran material  for  the  following  reasons.  Curran, 
in  mentioning  these  specimens,  made  them  syn- 
types;  he  does  not  indicate  that  he  saw  West's 
type,  and  there  is  a  slight  possibility  that  he  did 
not.  Also,  West's  type  is  in  poor  condition, 
whereas  Curran' s  material  is  in  good  condition, 
and  a  lectotype  designated  from  it  would  be  sup- 
ported by  a  long  series  of  paralectotypes.  I  have 
given  West's  Colorado  "holotype"  the  status  of 
misassociated  paralectotype. 

The  lectotype  and  paralectotypes  are  in  ex- 
cellent condition  and  are  deposited  in  AMNH, 
FSCA,  and  CUIC.  These  specimens  definitely 
belong  to  P.  rufipennis  (except  West's  "holo- 
type"). 

In  Curran 's  1930  key,  he  separated  P.  rufi- 
pennis, as  I  have  now  defined  it,  into  four 
species.  The  characters  he  used  are:  presence 
or  absence  of  infrasquamals,  color  of  male  legs, 
color  of  scutellum,  and  presence  or  absence  of 
parafacial  hairs.  All  except  the  last  are  character 
states  which  vary  among  P.  rufipennis  adults. 

DIAGNOSIS. — Specimens  of  P.  rufipennis  are 
distinguished  from  those  of  all  other  species  of 
Ptilodexia  by  the  following  combination  of  char- 
acters: face  long  and  narrow,  totally  devoid  of 
parafacial  hairs  ventral  to  second  antennal  seg- 
ment; antennae  long,  width  of  plumosity  on  aris- 
ta more  than  twice  length  of  second  antennal 
segment;  abdomen  grayish  black  or  dark  brown, 
reddish  laterally  in  a  few  individuals;  extreme 
tip  of  abdomen  and  genitalia  yellowish;  legs  of 
females  and  often  those  of  males  pale  in  color; 


WILDER:  NEARCTIC  PTILODEXIA 


45 


ejaculatory  apodeme  with  bulb  in  shape  of  a 
shallow  cup. 

VARIATION. — This  species  shows  remarkable 
intraspecific  variation.  Northern  specimens 
(those  from  Canada,  Michigan,  Wisconsin, 
North  Dakota,  and  Montana)  are  dark  in  facial 
ground  color;  the  antennae  are  dark  in  most 
adults;  the  legs  of  males  are  varied,  ranging  from 
light  brown  to  almost  black;  and  the  length  of 
the  haustellum  is  almost  half  the  head  height. 
Specimens  from  Florida  and  Georgia  into  Texas 
are  light  in  facial  ground  color;  the  antennae  are 
rufotestaceous  and  the  legs  of  males  are  pale, 
just  as  in  females.  These  specimens  have  the 
haustellum  short,  about  0.3  times  the  head 
height. 

In  intermediate  geographical  areas,  there  is  a 
mixture  of  the  two  forms.  It  is  not  uncommon 
to  have  a  series  split  into  half  dark  individuals 
and  half  light.  There  is  no  striking  difference  in 
the  genitalia  of  members  of  these  two  forms,  and 
in  the  areas  of  intergradation,  intermediates  oc- 
cur. These  intermediates  are  dark  with  short 
mouthparts  or  light  with  longer  ones,  and  they 
are  few  in  number.  Many  of  the  intermediate 
areas  are  represented  by  one  or  two  specimens, 
and  it  is  possible  that  a  representative  sample  of 
these  flies  will  show  the  presence  of  a  smooth 
cline,  northern  areas  having  the  largest  percent- 
age of  dark  specimens,  southern  areas  the  larg- 
est proportion  of  light  ones,  and  the  intermediate 
areas  showing  a  north-south  gradient  of  light 
and  dark. 

MATERIAL  EXAMINED. — The  sample  of  Ptilo- 
dexia  rufipennis  includes  694  males  and  612  fe- 
males. 

DISTRIBUTION. — This  species  ranges  from 
Newfoundland  to  southern  Florida,  across  the 
continent  to  British  Columbia,  southeastern 
Washington,  and  central  Montana.  There  are 
scattered  records  in  central  Colorado,  New 
Mexico,  and  Texas. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  May  to  November.  However,  most  speci- 
mens have  been  collected  in  the  months  of  July, 
August,  and  September.  In  the  Midwest  and 
Canada,  records  for  the  months  of  July  and  Au- 
gust predominate,  while  in  the  Northeast  and 
Mid-Atlantic  states  most  are  collected  during 
August  and  September.  In  areas  of  heavy  col- 
lecting (Suffolk  County,  New  York),  specimens 
seem  to  occur  with  similar  frequency  throughout 
the  season.  The  small  sample  from  Florida 


shows  specimens  collected  in  May  and  again  in 
September,  October,  and  November,  perhaps 
indicating  a  bivoltine  situation.  In  nearby  Geor- 
gia the  records  show  specimens  taken  from  mid- 
May  to  mid- August  with  most  occurring  during 
June  and  July. 

Ptilodexia  rufipennis  is  predominately  a  low- 
land species.  A  few  specimens  have  been  col- 
lected in  mountainous  areas.  In  New  Mexico, 
specimens  were  collected  at  2,440  m  in  the 
White  Mountains;  specimens  from  Georgia  were 
taken  up  to  1,460  m  in  the  southern  Appalachi- 
ans; up  to  1,890  m  in  North  Carolina;  and  up  to 
1,830  m  in  Tennessee. 

Ptilodexia  rufipennis  adults  have  been  col- 
lected on  a  number  of  different  flowers,  includ- 
ing the  following  Compositae:  Solidago  triner- 
vata,  Achillea  Millefolium,  Eupatorium 
perfoliatum,  Chrysanthemum  leucanthemum, 
and  unidentified  Solidago  sp.,  Aster  sp.,  Cir- 
sium  sp.,  and  Anaphalis  sp.  They  have  also 
been  noted  visiting  Melilotus  alba  (Fabaceae), 
Daucus  carota  (Umbelliferae),  Ceanothus 
americanus,  and  unidentified  Ceanothus  sp. 
(Rhamnaceae),  and  Lonicera  sp.  (Caprifoli- 
aceae).  Most  of  these  flowers  are  yellow  or 
white. 

One  specimen  was  collected  from  a  female 
asilid  (Proctocanthus  philadelphicus  Macquart), 
which  had  presumably  captured  it  as  prey.  Other 
means  of  collecting  have  been  by  net  (sweeping 
foliage),  UV  light  trap,  aerial  light,  and  Malaise 
trap.  Ptilodexia  rufipennis  adults  have  been  col- 
lected in  a  cranberry  bog,  marshy  meadow, 
creek  bank,  maple-elm  floodplain  along  stream, 
and  in  a  maple-mixed-oak-hickory  forest. 

Ptilodexia  arida  (Curran) 

(Figures  89-93) 

Rhynchiodexia  arida  CURRAN,  1931:93.  WEST  (1950:110);  SA- 
BROSKY  AND  ARNAUD  (1965:988);  COLE  (1969:543).  [LEC- 
TOTYPE  (here  designated),  a  female,  deposited  in  CUIC, 
labeled,  "Pinnaleno  Mts.,  Ft.  Grant,  Ariz.'VR.  C.  Shan- 
non July  17,  1917'VAltitude  7000  ft. "/"Wing  Slide,  Cornell 
U.,  Lot.  919,  Sub.  120,  S.  S.  West"/"?  Holotype  Rhyn- 
chiodexia arida  Curran'V'Holotype,  Cornell  U.,  No. 
1937"/"Cornell  U.,  Lot.  922,  Sub.  39'VLectotype  Rhyn- 
chiodexia arida  Curran  designated  by  D.  Wilder  1976."] 

TAXONOMIC  NOTES. — Curran  introduced  the 
name  into  the  literature  in  his  1930  key.  West, 
however,  was  the  one  to  segregate  and  name  the 
species.  West's  "holotype,"  a  female,  has  been 
designated  as  lectotype,  even  though  it  bears  a 
holotype  label  with  Curran  as  author.  A  speci- 


46 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


FIGURES  89-93.    Ptilodexia  arida  (Curran).  Fig.  89.  Genitalia  of  male,  posterior  view.   Fig.  90.  Ejaculatory  apodeme.   Fig. 
91.  Genitalia  of  male,  lateral  view.  Fig.  92.  Head  of  male,  lateral  view.   Fig.  93.  Geographical  distribution. 


men  from  Manitoba  has  been  designated  as 
paralectotype.  This  specimen  has  an  undated 
Curran  determination  label,  and  I  think  that  he 
compared  the  specimen  with  West's  type,  felt 
that  they  were  the  same,  and  consulted  both 
when  writing  the  1930  key. 

West's  type  is  a  female  in  poor  condition,  but 
the  Manitoba  male  is  in  good  condition.  The 
Manitoba  specimen  is  misassociated,  actually 
belonging  to  P.  major. 

DIAGNOSIS. — Ptilodexia  arida  adults  are 
unique  in  the  following  combination  of  charac- 
ters: broad,  square  face,  no  parafacial  hairs  ven- 
tral to  second  antennal  segment;  plumosity  on 
arista  long,  up  to  twice  length  of  second  antennal 
segment;  legs  of  females  pale  in  color;  abdomen 
of  males  and  many  females  reddish  laterally. 

MATERIAL  EXAMINED. — Sixty-four  males  and 
33  females  were  examined. 

DISTRIBUTION. — Ptilodexia  arida  ranges  from 
Veracruz  in  Mexico  into  the  mountains  of  Ari- 
zona and  Utah.  There  is  one  record  from  Jef- 
ferson County,  Idaho. 


BIOLOGICAL  NOTES. — The  flight  period  is 
from  March  to  October,  the  majority  of  speci- 
mens being  captured  in  July  and  August.  There 
is  only  one  March  record  and  none  for  the 
months  of  April  and  May.  One  specimen  from 
Morelos  in  Mexico  was  collected  in  November. 

This  species  inhabits  a  mountainous  region. 
Specimens  have  all  been  collected  in  the  moun- 
tains, some  at  elevations  up  to  2,560  m.  One 
exception  is  a  male  collected  in  Jefferson  Coun- 
ty, Idaho,  an  area  of  extensive  lava  fields.  Other 
specimens  have  been  collected  by  special  de- 
vices, such  as  UV  light  and  Malaise  traps.  One 
female  was  collected  on  Ceanothus  (Rhamna- 
ceae),  and  four  males  were  swept  from  Ipomoea 
(Convolvulaceae).  There  are  five  reared  speci- 
mens; however,  they  lack  data.  They  are  pinned 
with  their  hosts,  small  scarabaeid  larvae. 

Ptilodexia  ponderosa  (Curran) 

(Figure  6) 

Rhynchiodexia  ponderosa  CURRAN,  1930:93.  WEST  (1950:110); 
SABROSKY  AND  ARNAUD  (1965:989).  [LECTOTYPE,  female, 


WILDER:  NEARCTIC  PT1LODEXIA 


47 


FIGURES  94-98.    Ptilodexia  major  (Bigot).   Fig.  94.  Genitalia  of  male,  posterior  view.   Fig.  95.  Ejaculatory  apodeme.   Fig. 
96.  Genitalia  of  male,  lateral  view.  Fig.  97.  Head  of  male,  lateral  view.   Fig.  98.  Geographical  distribution. 


deposited  in  CUIC,  labeled,  "Chokoloskee,  Fla.'VWing 
Slide,  Cornell  U.  Lot.  919,  Sub.  129,  L.  S.  West"/"  9  Ho- 
lotype  Rhynchiodexia  ponderosa  Curran"/"Holotype  Cor- 
nell U.  No.  1936"/"Cornell  U.  Lot.  922,  Sub.  38'VLec- 
totype  Rhynchiodexia  ponderosa  Curran."] 

TAXONOMIC  NOTES. — This  species  was  orig- 
inally segregated  and  named  by  West,  but  the 
name  was  introduced  into  the  literature  by  Cur- 
ran  and  dates  from  his  1930  key.  The  unique 
specimen  was  not  designated  as  a  holotype. 
Since  Curran  did  not  mention  the  number  of 
specimens  which  he  had,  even  though  there  is 
only  one,  it  must  be  designated  lectotype.  The 
specimen  is  in  excellent  condition. 

DIAGNOSIS. — Since  this  species  is  represented 
by  one  specimen,  it  is  difficult  to  diagnose.  The 
P.  ponderosa  specimen  can  be  distinguished 
from  members  of  all  other  related  species  by  the 
following  combination  of  characters.  Size  large, 
color  light  reddish  brown;  face  wide;  parafacial 
hairs  small,  brown,  inserted  only  on  anterior  half 
of  parafacial;  vibrissal  angle  of  head  quite  pro- 
nounced; epistome  protrudes  noticeably;  third 
antennal  segment  long;  length  of  plumosity  1.6 
times  length  of  second  antennal  segment;  haus- 


tellum  length  0.6  times  head  height;  legs  uni- 
formly testaceous. 

DISTRIBUTION. — The  lectotype  was  collected 
in  extreme  southern  Florida  and  almost  certainly 
represents  a  West  Indian  species.  It  more 
closely  resembles  specimens  from  these  islands 
(most  of  which  have  small,  pale,  parafacial 
hairs,  long  plumosity  on  the  arista,  and  a  me- 
dium-long haustellum)  than  those  from  the 
United  States.  It  differs  from  the  specimens  I 
have  seen  from  the  islands  in  being  much  larger 
and  having  uniformly  pale  legs. 

major  Group 
Ptilodexia  major  (Bigot) 

(Figures  94-98) 

Rhamphinina  major  BIGOT  1888:265.  WULP  ( 1891:228);  GIG- 
Lio-Tos  (1894:60);  BRAUER  (1899:509);  ALDRICH  (1905:504); 
SABROSKY  AND  ARNAUD  (1965:989);  COLE  (1969:543);  Gui- 
MARAES  (1971:33).  [LECTOTYPE  (here  designated),  male, 
deposited  in  BMNH,  labeled,  "Lectotype" /"Rhamphinina 
major  Bigot'VMexico.  Ex.  coll.  Bigot.  Pres.  by  G.  H.  Ver- 
rall,  Oct.  1904.  1 904-274' V" Lectotype  Rhamphinina  major 
Bigot  designated  by  D.  Wilder  1975. "J 

Rhynchiodexia  tincticornis  BIGOT,  1888:266.  WULP  (1891:228); 
BRAUER  (1899:509);  ALDRICH  (1905:504);  GUIMARAES 


48 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


(1971:34).  [LECTOTYPE  (here  designated),  male,  deposited 
in  BMNH,  labeled,  "Lectotype" /"Rhynchiodexia  tincti- 
cornis  Bigot"/" Mexico.  Ex.  coll.  Bigot.  Pres.  by  G.  H. 
Verrall,  Oct.  1904.  1904-274"/Bigot's  original  inked  label/ 
Austen's  penciled  note  regarding  Bigot's  label/" Lectotype 
Rhynchiodexia  tincticornis  Bigot  designated  by  D.  Wilder 
1975."] 

Dexiosoma  fumipennis  BIGOT,  1888:270.  WULP  (1891:230); 
BRAUER  (1899:508);  ALDRICH  (1905:504);  GUIMARAES 
(1971:34).  [HoLOTYPE,  female,  deposited  in  BMNH,  la- 
beled ,  "Ho\otype" /"Rhynchiodexia  fumipennis  "/" Mexico. 
Ex.  coll.  Bigot.  Pres.  by  G.  H.  Verrall,  Oct.  1904.  1904- 
274"/2  folded  labels.]  NEW  SYNONYMY 

Rhynchodexia  fraterna  WULP,  1891:229.  GiGLio-Tos  (1894:61); 
ALDRICH  (1905:499);  THOMPSON  (1963:517);  GUIMARAES 
(1971:33).  [LECTOTYPE  (here  designated),  male,  deposited 
in  BMNH,  labeled,  "Lectotype'T'Cuernavaca,  Morelos. 
June  H.  H.  S."/"d"/"B.  C.  A.  Dipt.  II  Rhynchodexia  fra- 
terna, v.d.W. "/"Central  America  Pres.  by  F.  D.  Godman, 
O.  Salvin.  1903-172"/"Lectotype  Rhynchodexia  fraterna 
Wulp,  desig.  by  D.  Wilder  1975."]  NEW  SYNONYMY 

Rhynchodexia  omissa  WULP,  1891:235.  ALDRICH  (1905:499); 
GUIMARAES  (1971:33).  [LECTOTYPE  (here  designated), 
male,  deposited  in  BMNH,  labeled,  " Lectotype"/" N.  Yu- 
catan, Gaumer"/"cT'/"B.  C.  A.  Dipt.  II  Rhynchodexia  om- 
issa v.d.W. "/"Central  America  Pres.  by  F.  D.  Godman,  O. 
Salvin.  1903-172"/"Lectotype  Rhynchodexia  omissa  Wulp 
designated  by  D.  Wilder  1975."]  NEW  SYNONYMY 

Ptilodexia  harpasa  (partim):  DAVIS  (1919:83).  (misidentifica- 
tion) 

Rhynchiodexia  arida  (partim):  CURRAN  (1930:93).  (misidenti- 
fication) 

TAXONOMIC  NOTES. — The  confusion  and  syn- 
onymy of  this  species  in  the  literature  is  under- 
standable. It  is  quite  variable  in  color  charac- 
teristics and  size  of  its  members.  It  also  has  a 
broad  geographical  range.  Bigot  proposed  three 
names  for  P.  major  and  Wulp  two. 

The  syntype  series  for  P.  major  consists  of 
five  males.  Two  of  these  are  misassociated,  one 
resembling  specimens  of  P.  conjuncta  but  with 
a  shorter  haustellum,  the  other  resembling  spec- 
imens of  P.  scutellata  but  with  much  hairier 
parafacials.  The  other  three  are  conspecific.  The 
specimen  chosen  as  a  lectotype  has  the  upper 
parafacials  a  little  hairier  than  the  others,  and  its 
facial  tomentum  has  a  slightly  yellowish  cast. 
The  series  is  in  poor  condition,  the  lectotype 
having  only  one  leg  and  one  antenna;  most  of 
the  setae  are  broken. 

Associated  with  the  syntype  series  is  a  pen- 
ciled label  in  Austen's  handwriting.  It  mentions 
that  Brauer  (1899:509)  felt  the  species  belonged 
to  Ptilodexia.  He  continued,  however,  saying 
that  he  (Austen)  felt  that  the  series  was  com- 
prised of  representatives  of  three  distinct  gen- 
era, only  one  specimen  belonging  to  Ptilodexia. 
The  specimen  which  Austen  felt  was  a  Ptilo- 


dexia is  the  misassociated  type  resembling  spec- 
imens of  P.  conjuncta.  His  reasoning  is  not 
clear,  but  perhaps  he  failed  to  see  the  minute 
parafacial  hairs  on  true  P.  major  members  and 
therefore  felt  it  belonged  in  another  genus.  The 
type  which  resembles  adults  of  P.  scutellata 
could  understandably  be  considered  as  repre- 
senting a  different  genus  by  a  worker  who  be- 
lieved in  restricted  genera. 

Bigot  stated  the  type-locality  as  Mexico  and 
Washington  Territory.  All  five  syntypes  have  la- 
bels reading  "Mexico." 

Both  of  the  cotypes  of  Rhynchiodexia  tincti- 
cornis are  in  poor  condition.  The  specimen  cho- 
sen as  lectotype  is  oily,  the  tomentum  therefore 
obscured.  Wulp  (1891)  redescribed  the  species 
and  restricted  the  type-locality  to  Paso  del 
Macho,  possibly  taking  the  name  of  the  locality 
from  another  Bigot  label.  Since  this  locality  is 
within  the  range  of  the  species,  the  restriction 
is  accepted.  Wulp  stated  that  he  had  a  female 
from  Costa  Rica.  It  is  possible,  but  unlikely,  that 
the  species  ranges  that  far  south.  Brauer  (1899) 
placed  P.  major  in  Ptilodexia  and  listed  P.  tinc- 
ticornis as  a  synonym. 

Bigot  described  R.  fraterna  in  1891  from  21 
males  and  5  females.  Of  the  four  male  specimens 
which  I  had  the  opportunity  to  see,  one  was 
misassociated,  the  other  three  belonged  to  P. 
major.  Wulp  himself  stated  after  his  description 
that  three  of  his  largest  specimens  "agree  fully 
with  a  typical  example  of  Rhamphinina  major,'" 
and  he  then  cited  a  few  minor  color  differences. 
He  also  felt  that  some  of  the  smaller  specimens 
"bear  a  striking  resemblance"  to  P.  fumipennis 
adults.  The  lectotype,  a  male  from  Cuernavaca, 
is  in  good  condition. 

Rhynchiodexia  omissa  was  described  from 
three  male  and  two  female  cotypes.  The  male 
which  I  saw  and  designated  as  lectotype  belongs 
to  P.  major.  The  specimen  was  collected  in 
northern  Yucatan.  It  is  in  excellent  condition 
and  is  similar  in  color  to  the  lectotype  of  P. 
tincticornis.  Guimaraes  (1971)  was  the  first  to 
place  omissa  in  Ptilodexia. 

Another  name  must  be  considered  while  dis- 
cussing P.  major.  This  is  Rhamphinina  dubia 
Bigot,  the  type-species  of  that  genus.  Bigot  de- 
scribed R.  dubia  from  two  females  labeled  only 
"Mexico."  The  two  specimens  now  in  the  Brit- 
ish Museum  are,  however,  a  male  and  a  female. 
The  male  belongs  to  P.  major  and  the  female  to 
Macrometopa  calogaster,  the  latter  placed  by 


WILDER:  NEARCTIC  PTILODEXIA 


49 


Austen.  Townsend  (1931:100)  spoke  of  a  female 
holotype;  however,  this  cannot  be  considered  a 
lectotype  designation  of  the  type-series  accord- 
ing to  Crosskey  (in  litt.).  I  have  designated  the 
female  of  the  original  material  as  the  lectotype, 
giving  the  male  the  status  of  misassociated  para- 
lectotype.  Had  the  other  specimen  ( 6 ,  P.  major) 
been  chosen  as  the  lectotype,  the  name  Rham- 
phinina  (1885)  would  have  priority  over  Ptilo- 
dexia  (1889)  and  our  commonly  used  name 
would  have  to  be  changed.  Macrometopa  now 
becomes  a  synonym  of  Rhamphinina. 

DIAGNOSIS. — Ptilodexia  major  exhibits  much 
variation  throughout  its  range,  and  after  a  more 
thorough  sample  is  available,  may  be  found  to 
consist  of  more  than  one  species.  It  is  most  dif- 
ficult to  diagnose,  and  specimens  can  be  more 
easily  identified  by  first  eliminating  other 
species. 

Thorax  dark,  the  notum  distinctly  striped; 
parafacial  hairs  small,  pale,  inserted  only  on  an- 
terior half  of  parafacial,  ventrally  on  face  to  apex 
of  antennae;  they  are  light  brown,  medium 
length,  and/or  scattered  on  parafacial  in  few  in- 
dividuals; haustellum  length  varied,  from  0.55  to 
0.65  times  head  height,  in  some  small  specimens 
it  is  slightly  longer;  palpi  longer  than  half  haus- 
tellum length  in  most  individuals;  length  of  plu- 
mosity  on  arista  more  than  twice  length  of  sec- 
ond antennal  segment;  abdomen  reddish  with 
dark  median  stripe  in  most  males,  entirely  dark 
or  dark  with  fourth  tergite  reddish  in  a  few. 

VARIATION. — Ptilodexia  major  exhibits  more 
variability  than  do  most  Ptilodexia.  A  summary 
of  this  variation  is  given  by  Wilder  (1976). 

There  are  some  specimens,  notably  those 
from  Texas,  New  Mexico,  Colorado,  and  Ne- 
braska, which  seem  to  fall  somewhere  between 
specimens  of  P.  incerta  and  P.  major,  two 
species  which  are,  from  available  records,  allo- 
patric.  It  is  possible  that  these  represent  an  as 
yet  unrecognized  species  and  that  true  P.  major 
is  a  more  southern  species.  Most  of  the  char- 
acters involved,  however,  are  vague  and  in  other 
species  are  part  of  normal  intraspecific  varia- 
tion. This  problem  may  be  solved  when  a  larger 
sample  is  available,  thereby  allowing  the  use  of 
statistical  techniques.  I  have  placed  these  spec- 
imens in  P.  major  because  it  is  with  members 
of  this  species  that  they  share  the  most  character 
states. 

Most  Nearctic  Ptilodexia  do  not  have  ranges 
which  extend  into  neotropical  Mexico  (P.  con- 


junct a  is  an  exception).  Specimens  of  P.  major 
from  Chiapas,  Campeche,  and  Yucatan,  how- 
ever, differ  very  little  from  those  from  Arizona, 
at  least  in  characters  which  easily  separate 
adults  or  other  species  in  the  genus.  I  feel  jus- 
tified in  placing  all  these  specimens  in  P.  major, 
although  when  more  Mexican  material  is  avail- 
able for  study,  it  may  show  that  more  than  one 
species  is  involved. 

MATERIAL  EXAMINED. — One  hundred  fifty- 
two  males  and  77  females  of  P.  major  were  ex- 
amined. 

DISTRIBUTION. — This  species  ranges  from 
Nebraska  southward  through  Colorado,  Kansas, 
Arizona,  New  Mexico,  and  Texas  into  southern 
Mexico. 

BIOLOGICAL  NOTES. — The  flight  period  ranges 
from  April  to  October,  with  the  main  activity  in 
July  and  August  in  most  parts  of  the  range. 
Members  of  Texas  populations  exhibit  two  pe- 
riods of  activity,  one  in  April  and  May,  and 
another  in  September  and  October.  Specimens 
reared  indoors  appear  to  emerge  much  earlier, 
in  December,  February,  and  March. 

Specimens  have  been  taken  at  both  low  and 
high  elevations,  the  highest  being  2,130  m  in 
Coahuila,  Mexico.  Methods  used  to  collect  P. 
major  adults  are  sweeping,  UV  light  trap,  Ma- 
laise trap,  and  rearing.  Flowers  from  which  P. 
major  adults  have  been  captured  include:  Se- 
necio  sp.,  Baccharis  glutinosa  (Compositae); 
Lepidium  thurberi  (Cruciferae);  Croton  sp.  (Eu- 
phorbiaceae);  Medicago  saliva.  Dale  a  albiflora 
(Fabaceae);  Marrubium  vulgare  (Labiatae); 
Gossypium  sp.  (Malvaceae),  and  Tamarix  sp. 
(Tamaricaceae).  Unlike  other  Ptilodexia,  com- 
posites are  not  the  group  of  flowers  most  fre- 
quently visited. 

Several  specimens  among  the  material  exam- 
ined have  been  reared  and  are  associated  with 
skins  of  host  larvae.  Unfortunately,  these  skins 
cannot  be  identified.  Host  data  on  some  speci- 
mens list  the  host  as  "May  beetle"  and  Phyl- 
lophagafarcta.  David  (1919)  mentions  Phylloph- 
aga  grubs  collected  on  29  April,  at  Austin, 
Texas: 


Ptilodexia  larvae  were  first  observed  from  May  18  to  June 
18.  In  these  cases,  adult  flies  issued  between  June  11  and 
July  10,  the  puparium  stage  varying  from  20-32  days.  Grubs 
collected  in  New  Mexico  May  10  and  24  showed  parasitism 
from  May  20  to  July  7,  while  larvae  from  the  same  locality 
collected  on  September  7  and  confined  in  indoor  cages 
showed  parasitic  larvae  from  October  7  to  February  23. 


50 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


FIGURES  99-103.    Ptilodexia  incerta  (West).  Fig.  99.  Genitalia  of  male,  posterior  view.  Fig.  100.  Ejaculatory  apodeme.  Fig. 
101.  Genitalia  of  male,  lateral  view.  Fig.  102.  Head  of  male,  lateral  view.  Fig.  103.  Geographical  distribution. 


Aldrich  called  these  New  Mexico  and  Texas 
specimens  Ptilodexia  harpasa,  but  they  are  typ- 
ical of  P.  major  adults.  Davis  also  reports  mem- 
bers of  this  species  being  reared  from  grubs  of 
Aphonus  pryiformis  collected  in  Las  Vegas  and 
Caffrey,  New  Mexico. 

Ptilodexia  incerta  West 
(Figures  99-103) 

Ptilodexia  incerta  WEST,  1925:131.  LEONARD  (1928:822); 
CURRAN  (1930:93);  SABROSKY  AND  ARNAUD  (1965:989). 
[HoLOTYPE,  male,  deposited  in  CUIC,  labeled,  "River- 
head,  L.I.,  N.Y.  6-30-2 1'VHolotype  Ptilodexia  incerta 
West  cTT-Holotype  Cornell  U.  No.  1873'VCornell  U. 
Lot.  922,  Sub.  45"] 

Ptilodexia  proximo  WEST,  1925:133.  LEONARD  (1928:822); 
CURRAN  (1930:93);  WEST  (1950:pl.  I,  fig.  4);  SABROSKY  AND 
ARNAUD  (1965:989).  [HOLOTYPE,  male,  deposited  in  CUIC, 
labeled,  "Riverhead,  L.I.,  N.Y.'VWm.  T.  Davis,  Aug.  5, 
1917'VWing  Slide,  Cornell  U.  Lot.  919,  Sub.  121,  L.  S. 
West'V'Holotype  Ptilodexia  proximo  West  d'V'Holotype 
Cornell  U.  No.  1877"/"Cornell  U.  Lot  922,  Sub.  47"]  NEW 
SYNONYMY 

Rhynchodexia  elevata  WEST,  1925:135.  LEONARD  (1928:821); 
SABROSKY  AND  ARNAUD  (1965:988).  (HOLOTYPE,  male,  de- 
posited in  CUIC,  labeled,  "Riverhead,  L.I.,  N.Y.,  Aug.  20, 
1922"/"Wing  Slide  Cornell  U.  Lot.  919,  Sub.  123,  L.  S. 


West'V'Holotype  Rhynchodexia  elevata  West  cJ"/"Holo- 
type  Cornell  U.  No.  1871"rCornell  U.  Lot.  922,  Sub.  36."] 
NEW  SYNONYMY 

Ptilodexia  harpasa  (partim):  JOHNSON  (1925b:208).  (misiden- 
tification) 

Rhynchiodexia  levata  West:  CURRAN  (1930:93).  (lapsus  cala- 
mus) 

TAXONOMIC  NOTES. — West  described  P.  in- 
certa and  its  two  synonyms  in  the  same  paper. 
The  type-localities  of  all  three  are  Riverhead, 
Long  Island.  None  of  his  holotypes,  all  males, 
show  any  significant  differences.  The  type  of 
P.  proxima  is  large,  that  of  P.  elevata  is  small, 
while  the  type  of  P.  incerta  is  intermediate  in 
size. 

West  placed  P.  elevata  in  the  genus  Rhyn- 
chodexia because  he  thought  that  it  lacked  para- 
facial  hairs.  These  hairs,  however,  are  present 
and  are  similar  in  size,  color,  and  number  to 
those  of  the  other  two  species.  He  also  felt  that 
the'/*,  elevata  adult  differed  in  having  the  vi- 
brissae  inserted  far  above  the  oral  margin, 
another  characteristic  which  is  the  same  in  the 
other  two  species. 


WILDER:  NEARCTIC  PT1LODEXIA 


51 


West  stated  no  diagnostic  difference  between 
the  P.  incerta  and  P.  proxima  males,  and  the 
descriptions  of  the  two  are  nearly  identical. 

The  types  of  these  three  species  names  are  in 
excellent  condition,  with  the  possible  exception 
of  the  holotype  of  P.  incerta,  which  is  dusty. 
They  are  all  deposited  in  CUIC.  The  three  are 
unquestionably  conspecific. 

DIAGNOSIS. — Adults  of  P.  incerta  can  be  dis- 
tinguished from  those  of  other  species  of  the  ge- 
nus overlapping  in  range  by  the  following  com- 
bination of  character  states:  parafacial  hairs  tiny 
and  pale  (rarely  brown),  present  only  on  ex- 
treme upper  anterior  portion  of  parafacial; 
length  of  third  antennal  segment  about  1.5  times 
length  of  second;  length  of  plumosity  on  arista 
twice  length  of  second  antennal  segment.  Haus- 
tellum  length  0.5  to  0.6  times  head  height;  palpi, 
broad  in  many  individuals,  equal  to  or  less  than 
half  haustellum  length.  Thorax  distinctly  striped; 
in  many  specimens  there  are  few  or  no  infra- 
squamal  setulae;  in  most  abdomen  reddish  with 
a  median  longitudinal  stripe. 

MATERIAL  EXAMINED. — Specimens  exam- 
ined include  105  males  and  32  females. 

DISTRIBUTION. — This  species  occurs  through- 
out the  eastern  U.S.,  west  to  about  the  100th 
meridian. 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  May  to  September,  with  most  specimens 
being  collected  during  July.  In  certain  areas,  es- 
pecially in  the  Southeast,  specimens  have  been 
taken  in  every  month  from  10  May  to  26  Sep- 
tember. 

Associated  data  show  that  P.  incerta  adults 
have  been  collected  while  at  blooms  of  Cirsium 
sp.  and  Solidago  sp.,  both  Compositae.  They 
have  also  been  taken  on  high  prairie  in  Hennepin 
County,  Minnesota.  A  male  was  taken  from  a 
phymatid,  which  had  presumably  captured  it  as 
prey.  Records  show  that  P.  incerta  adults  have 
only  been  collected  at  low  elevations. 

Ptilodexia  maculata,  new  species 

(Figures  104-108) 

TYPE-LOCALITY. — The  holotype  was  collected  14  miles  [ca. 
22.5  km]  north  of  Silver  City,  Grant  County,  New  Mexico. 

TYPE-SPECIMENS. — The  holotype,  a  male,  is  deposited  in 
MSUC,  the  allotype,  from  Show-Low  Arizona,  in  USNM. 
Complete  data  from  these  specimens  and  the  12  male  and  10 
female  paratypes  are  listed  below. 

DIAGNOSIS. — Ptilodexia  maculata  is  a  distinc- 
tive species,  its  members  differing  from  those  of 


its  congeners  by  the  following  combination  of 
character  states.  Reddish  brown;  face  wide,  an- 
teroventral  angle  not  strongly  protruding;  para- 
facial hairs  small,  pale,  occuring  only  along  an- 
terior edge  of  parafacial;  oral  margin  only 
slightly  protruding;  infrasquamal  setulae  lack- 
ing; haustellum  length  0.4  to  0.5  times  head 
height.  The  legs  of  males  and  females  are  pale, 
the  femora  with  striking  dark  brown  apicoven- 
tral  patches. 

DESCRIPTION. — Male:  Total  body  length  16 
mm,  brownish  with  median  abdominal  stripe. 
Head  height  3.5  mm;  profrons  0.3  times  head 
length.  Facial  tomentum  pale  gold,  shining,  fine, 
not  obscuring  integumental  color;  parafrontal 
ha"irs  long,  dark,  sparse;  parafacial  hairs  small, 
pale,  inserted  only  along  anterior  edge  of  para- 
facial and  not  ventral  to  antennal  apex.  Width 
of  frontal  vitta  at  base  of  antenna  0.55  mm,  at 
narrowest  part  0.25  mm;  carina  well  developed, 
long,  wide;  facial  cavities  slightly  darkened. 
Height  of  gena  0.35  times  head  height.  One  pair 
of  ocellar  setae,  2  pairs  of  postocellars  subequal 
to  ocellars,  1  pair  of  shorter  postverticals,  1  pair 
of  long  inner  verticals,  and  1  pair  of  outer  ver- 
ticals subequal  to  postorbitals.  Fifteen  pairs  of 
postorbital  setae  not  interspersed  with  smaller 
hairs,  those  closest  to  ocellar  triangle  long,  be- 
coming smaller  with  distance  from  ocellar  tri- 
angle, curved  only  slightly  anterad;  between 
postorbitals  and  the  dense  whitish  occipital  hair 
are  a  few  scattered  hairs.  Twelve  pairs  of  frontal 
setae;  one  pair  of  oral  vibrissae  0.25  mm  from 
oral  margin,  the  depression  between  them  0.4 
mm  wide;  12  additional  oral  setae,  1  above  vi- 
brissae fairly  strong,  peristomals  subequal  to 
each  other.  Epistome  only  slightly  protruding, 
its  width  1.3  times  length  of  first  two  antennal 
segments.  Haustellum  broad,  brown,  0.4  times 
head  height;  palpi  testaceous,  0.7  times  haustel- 
lum length.  Antennae  unique;  second  segment 
light  brown,  testaceous  apically;  third  segment 
testaceous  on  basal  half,  light  brown  apically; 
arista  and  plumosity  pale  brown.  Third  segment 
of  antenna  slightly  narrowed  apically,  1.3  times 
length  of  second  segment;  length  of  plumosity 
on  arista  1.5  times  length  of  second  antennal 
segment.  Thorax:  Mesonotum  rufotestaceous, 
brown  posteriorly,  subshining;  tomentum  beige, 
striping  indistinct.  Integument  of  humeri  and 
postalar  calli  rufotestaceous.  Six  pairs  of  strong 
humeral  setae,  1  posthumeral,  1  presutural,  and 
2  pairs  of  discal  scutellars.  Pleura  rufotestaceous 


52 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


107 


FIGURES  104-108.    Ptilodexia  maculata,  n.sp.  Fig.  104.  Genitalia  of  male,  posterior  view.  Fig.  105.  Ejaculatory  apodeme. 
Fig.  106.  Genitalia  of  male,  lateral  view.  Fig.  107.  Head  of  male,  lateral  view.  Fig.  108.  Geographical  distribution. 


marked  with  brown,  tomentum  beige,  translu- 
cent; infrasquamal  setulae  absent;  squamae  and 
wings  pale  testaceous;  epaulet  light  brown,  ba- 
sicosta  testaceous,  mediotergite  tomentose, 
subshining.  Legs  rufotestaceous,  tibiae  dark- 
ened basally  and  apically,  coxae  and  trochanters 
marked  with  dark  brown;  femora  with  dark 
brown,  strongly  contrasting  ventral  patches  on 
apical  half,  bases  of  setae  inserted  on  dark 
patches  pale  in  color,  pale  setal  bases  on  mac- 
ulae of  anterior  femur  coalesced  into  pale  stripe; 
tarsi  black.  Anterior  femur  with  12  dorsal  setae, 
8  posterodorsals,  and  13  posteroventrals,  tibia 
with  a  distinct  dark  dorsal  line.  Middle  femur 
with  2  anterior  setae  in  1  row.  Posterior  femur 
with  a  row  of  7  anterodorsal  setae,  9  anteroven- 
trals,  6  posteroventrals,  and  1  anterior  seta;  pos- 
terior tibia  curved.  Abdomen  rufotestaceous 
with  dark  brown,  narrow,  median  stripe;  tomen- 
tum whitish,  brown  above  median  stripe.  First 
syntergite  with  3  pairs  of  lateral  marginal  setae. 
Second  tergite  with  3  pairs  of  median  discals,  12 
dorsal  and  lateral  marginals,  and  4  pairs  of  lat- 
eral discal  setae.  Ventral  margins  of  tergite 


marked  with  brown.  Genitalia:  Eighth  tergite 
with  numerous  fine,  dark  setae.  Ninth  tergite 
with  many  long,  dark  setae,  lateral  swelling 
small.  Surstylus  huge,  lateral  angle  smoothly 
rounded,  depression  fairly  deep,  with  only  fine 
hairs;  in  profile,  smoothly  rounded  with  slight 
ventral  swelling.  Cerci  elongate,  lateral  lobes 
laterally  curved  strongly  anterad,  mesal  margins 
slightly  concave;  height  of  arms  slightly  greater 
than  that  of  lobes,  arms  not  reaching  ventral 
margins  of  surstyli,  in  profile,  slightly  bulging, 
tips  large,  round.  Ejaculatory  apodeme  with 
bulb  and  stem  distinct,  the  latter  slightly  longer 
and  curved.  Bulb  widely  open  on  one  side  and 
top;  opposite  side  strongly  decumbent.  Female: 
Similar  to  male  except  for  usual  sexual  differ- 
ences and  the  following.  Width  of  frontal  vitta 
at  narrowest  point  0.4  mm,  at  antennal  base  0.7 
mm.  Height  of  gena  0.35  times  head  height.  Ten 
pairs  of  short  postorbital  setae,  with  1  or  2  small 
hairs  interspersed;  7/8  frontal  setae.  Anterior  fe- 
mur with  9  posteroventrals,  7  posterodorsals, 
and  9  dorsal  setae.  Middle  femur  with  4  anterior 
setae  in  2  rows,  4  posteroventrals,  and  no  an- 


WILDER:  NEARCTIC  PTILODEX1A 


53 


FIGURES  109-113.    Ptilodexia  flavotessellata  (Walton).    Fig.  109.  Genitalia  of  male,  posterior  view.   Fig.  110.  Ejaculatory 
apodeme.   Fig.  1 1 1.  Genitalia  of  male,  lateral  view.   Fig.  1 12.  Head  of  male,  lateral  view.  Fig.  1 13.  Geographical  distribution. 


teroventrals.  Posterior  femur  with  no  anterior 
setae,  5  anterodorsals,  3  posteroventrals,  and  3 
anteroventrals.  Abdomen  rufous  with  vague 
dark-brown  median  stripe,  pollen  whitish,  light 
brown  around  median  marginal  setae.  First  syn- 
tergite  with  1  pair  of  strong  lateral  marginal  se- 
tae. Second  tergite  with  1  pair  of  lateral  margin- 
als, median  discals,  and  median  marginals.  Third 
tergite  with  1  pair  of  median  discals  and  5  dorsal 
and  lateral  marginal  setae. 

VARIATION. — Intraspecific  variation  is  slight. 
Total  body  length  ranges  from  10  to  16  mm.  The 
parafacial  hairs  are  brownish  in  some  speci- 
mens, but  they  are  small  in  all  adults.  The  area 
between  the  oral  vibrissae  and  the  oral  margin 
has  the  height  equal  to  width  in  some  specimens. 
The  integument  of  the  notum  in  some  individuals 
is  rufotestaceous  striped  with  dark  brown.  All 
facial  and  body  tomentum  are  shades  of  brown 
in  some  adults,  in  females,  quite  dark. 

MATERIAL  EXAMINED. — Thirteen  males  and  eleven  females 
were  examined  from  the  following  localities.  Arizona:  Corhise 
Co.:  Id,  Garden  Canyon,  Huachuca  Mts.,  30  July  1949,  W. 
Gertsch  (AMNH);  \<3,  Ramsey  Canyon,  15  miles  [ca.  24  km] 


S  of  Sierra  Vista,  6,000  ft  [ca.  1,830m],  1  July  1964,  Sternitzky 
(CNCI);  Id,  Rucker  Canyon,  Chiricahua  Mts.,  22  Aug.  1965, 
G.  Wallace  (UCRC).  Navajo  Co.:  19,  Show-Low,  24  July 
1956,  Butler  (UAIC).  Santa  Cruz  Co.:  19,  Madera  Canyon, 
4,880  ft  [ca.  1,490  m],  15  June  1965,  D.  Harrington,  19,  25 
July,  Id,  6  Aug.  1961,  at  light,  G.  Nelson,  19,  1 1  Aug.  1965, 
D.  Harrington,  29  9,  12  Aug.,  Id,  15  Aug.,  Id,  17  Aug.  1972, 
D.  Knull,  Idl9,  26  Aug.  1965,  D.  Harrington,  Id,  1  Sep., 
Id39  9,  7  Sep.,  19,  15  Sep.  (UCDC,  OSUC).  New  Mexico: 
Catron  Co.:  2dd,  Mogollon  Mts.,  29  Aug.  1951  (CASC). 
Grant  Co. :  1  d ,  14  miles  [ca.  22.5  km]  N  of  Silver  City,  8  July 
1961,  G.  Eickwort  (MSUC).  Hidalgo  Co.:  Id,  Guadalupe 
Canyon,  7  Aug.  1967,  J.  Smith  (UCRC). 

BIOLOGICAL  NOTES. — The  flight  period  lasts 
from  July  to  September.  Adults  of  this  species 
have  been  collected  at  lights.  They  inhabit 
mountainous  areas  and  have  been  taken  at  ele- 
vations from  1,490  m  to  1,830  m. 

DERIVATION  OF  NAME. — The  name  of  this 
species  refers  to  the  striking  femoral  maculae  on 
its  members. 

Ptilodexia  flavotessellata  (Walton) 

(Figures  109-113) 

Rhynchiodexia  flavotessellata   WALTON,    1914:176.   TOWN- 
SEND  (1915:23);  (1938:349);  SABROSKY  AND  ARNAUD 


54 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 


(1965:988);  COLE  (1969:543).  [HOLOTYPE,  female,  taken 
in  copula  and  pinned  with  its  mate,  deposited  in  USNM,  la- 
beled, "Koehler,  New  Mexico,  8-14'VWebster  No. 
7707"/"W.  R.  Walton  collector."] 

TAXONOMIC  NOTES. — Walton  stated  that  he 
described  this  species  from  four  specimens  tak- 
en at  Eagle  Tail  Mountain,  Colfax  County,  New 
Mexico.  He  named  a  female,  pinned  in  copula 
with  its  mate,  the  holotype.  These  specimens 
bear  a  label  which  reads  only,  "Koehler,  New 
Mexico"  as  the  locality;  however,  since  Walton 
himself  was  the  collector,  his  restriction  is  ac- 
cepted. There  are  more  specimens  than  the  de- 
scription records  with  identical  labels,  so  the 
three  males  which  he  indirectly  makes  paratypes 
cannot  be  segregated.  The  type  material  is  in  ex- 
cellent condition. 

Townsend  (1915)  erected  the  genus  Myocer- 
opsis  for  P.  flavotessellata.  He  believed  that 
specimens  of  Ptilodexia  had  the  haustellum  lon- 
ger than  0.75  times  the  head  height  and  that  P. 
flavotessellata,  its  members  with  the  haustellum 
length  0.5  times  the  head  height,  differed  suffi- 
ciently to  be  segregated  generically. 

DIAGNOSIS. — Specimens  of  P.  flavotessellata 
can  be  distinguished  from  those  of  other  North 
American  Ptilodexia  by  the  following  combi- 
nation of  character  states:  general  color  pale  or- 
ange-brown, tarsi  darker;  parafacial  hairs  pale 
to  brown,  of  medium  length,  sparse,  scattered 
randomly  on  parafacial;  third  antennal  segment 
not  much  longer  than  second;  length  of  plumos- 
ity  1.25  to  1.5  times  length  of  second  antennal 
segment;  haustellum  light  brown  in  color,  nar- 
row, about  0.5  times  head  height;  area  between 
oral  vibrissae  and  oral  margin  wider  than  high; 
epistome  protrudes  only  slightly. 

MATERIAL  EXAMINED. — Five  male  and  three 
female  specimens  were  examined. 

DISTRIBUTION. — Ptilodexia  flavotessellata 
ranges  from  northeastern  New  Mexico  north 
into  Colorado  and  Nebraska. 

BIOLOGICAL  NOTES. — The  collection  dates  on 
the  specimens  examined  range  from  22  June  in 
Boulder,  Colorado,  to  9  September  in  Nebraska. 
There  are  no  associated  ecological  data. 

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PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  2,  pp.  57-67,  6  figs.,  2  tables. 


March  2,  1979 


THE  SNAKE  EELS  (PISCES,  OPHICHTHIDAE)  OF  THE  HAWAIIAN 
ISLANDS,  WITH  THE  DESCRIPTION  OF  TWO  NEW  SPECIES 

By 
John  E.  McCosker 

Steinhart  Aquarium,  California  Academy  of  Sciences,  San  Francisco,  California  94118 


ABSTRACT:  The  ophichthid  eels  of  the  Hawaiian  Islands  (including  Johnston  and  the  Leeward  islands)  are 
reviewed;  included  are  species  new  to  Hawaii  and  extralimital  records  of  species  previously  considered  to  be 
endemic.  A  key  to  species  identification  is  provided.  Two  new  species  captured  in  deepwater  traps  off  Oahu  are 
described:  Muraenichthys  puhioilo,  subfamily  Myrophinae,  captured  at  275  m,  and  Ophichthus  kunaloa, 
subfamily  Ophichthinae,  captured  at  350  m.  Data  concerning  the  following  species  are  provided:  Schismorhyn- 
chus  labialis,  Muraenichthys  cookei,  M.  macropterus ,  Apterichtus  flavicaudus ,  Ichthyapus  vulturis,  Phaenomonas 
cooperae,  Callechelys  luteus,  Myrichthys  maculosus,  M.  bleekeri,  Cirrhimuraena  playfairii,  Brachysomophis  sau- 
ropsis,  B.  henshawi,  Phyttophichthus  xenodontus,  Ophichthus  polyophthalmus  and  ().  erabo.  Differences  in  ver- 
tebral number  of  populations  of  Myrichthys  maculosus  are  discussed  and  the  eastern  Pacific  nominal  species  M. 
xy stunts  (Jordan  &  Gilbert),  M.  tigrinus  Girard  and  M.  pantostigmius  Jordan  &  McGregor  are  placed  in  its 
synonymy.  The  endemism  (5  of  the  15  species)  of  the  Hawaiian  ophichthid  fauna  and  the  problems  of  populations 
and  species  differences  are  discussed. 


INTRODUCTION 

The  snake  eels,  family  Ophichthidae,  of  the 
Hawaiian  Islands  (including  Johnston  and  the 
Leeward  islands)  were  first  treated  by  Jordan 
and  Evermann  (1905)  and  subsequently  re- 
viewed by  Gosline  (1951)  and  Gosline  and  Brock 
(1960).  Recent  collections  by  the  George  Van- 
derbilt  Foundation,  John  E.  Randall  of  the  Bish- 
op Museum,  and  Thomas  A.  Clarke  of  the  Uni- 
versity of  Hawaii  have  added  important 
additional  specimens.  The  Hawaiian  Archipela- 
go contains  a  particularly  interesting  eel  fauna 
in  terms  of  its  abundance  and  the  range  of  dis- 
tributional conditions  which  exist,  including 
species  that  are  endemic  to  the  islands  as  well 
as  those  that  are  distributed  eastward  to  Austra- 
lia and  the  Red  Sea.  This,  while  recognizing  the 
dispersal  mechanism  allowed  by  the  leptoceph- 
alus  larva,  provides  an  intriguing  study  for  ma- 


rine zoogeographers.  Those  considerations,  as 
well  as  the  recent  capture  of  other  Hawaiian 
ophichthids  and  two  apparently  undescribed 
deepwater  species,  have  prompted  this  review. 

METHODS 

All  measurements  are  straight-line  (point  to 
point).  Standard  length,  trunk  length,  and  tail 
length  were  read  on  a  300-mm  ruler  with  0.5-mm 
gradations  and  were  recorded  to  the  nearest  0.5 
mm.  All  other  measurements  were  made  with 
dial  calipers  and  were  recorded  to  the  nearest 
0.1  mm.  Head  length  was  measured  from  the 
snout  tip  to  the  posterodorsal  margin  of  the  gill 
opening;  trunk  length  was  taken  from  the  end  of 
the  head  to  mid-anus;  body  depth  does  not  in- 
clude the  fin.  Vertebrae  (which  include  the  last 
centrum)  were  counted  from  radiographs. 

Comparisons  are  based  in  part  on  specimens 


[57] 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES.Vol.  42,  No.  2 


extralimital  to  the  Hawaiian  Islands  when  ma-  vomer  in  broad  bands;  snout  bluntly 

terial  was  insufficient.  Partial  synonymies  are  rounded Muraenichthys  schultzei 

provided  where  applicable  to  the  Hawaiian  Is-  4b.  Posterior  nostril  inside  mouth,  covered 

lands  taxa  and  where  new  synonymies  are  pro-  externally  by  a  flap;  teeth  uniserial  or 

posed.  biserial,  not  in  broad  bands;  snout  either 

Specimens  utilized  in  this  study  are  deposited  blunt  or  acute 5 

in  the  following  institutions:   Australian   Mu-  5a.  DFO  anterior  to  anus,  about  midway  to 

seum,  Sydney  (AMS);  Academy  of  Natural  Sci-  gill  openings  

ences  of  Philadelphia  (ANSP);  British  Museum  Muraenichthys  puhioilo  n.sp. 

(Natural  History)  (BMNH);  Bernice  P.  Bishop       5b.  DFO  above  or  behind  anus  6 

Museum  (BPBM);  California  Academy  of  Sci-  6a.  Snout  blunt;  DFO  above  or  slightly  be- 

ences  (CAS),  now  including  the  George  Van-  fore  anus Muraenichthys  cookei 

derbilt  Foundation  (GVF)  and  the  Stanford  Uni-  6b.  Snout     acute;     DFO     slightly     behind 

versity  collections  (SU);  Hawaii  Institute  of  anus Muraenichthys  gymnotus 

Marine  Biology  (HIMB);  Los  Angeles  County  7a.  Body  entire  finless;  coloration  either  uni- 

Museum  of  Natural  History  (LACM);  Scripps  form  or  darker  dorsally,  without  large 

Institution  of  Oceanography  (SIO);  University  spots  or  saddles  8 

of  Hawaii  (UH);  and  the  National  Museum  of  7b.  At  least  a  minute,  short,  dorsal  fin  pres- 

Natural  History  (USNM).  ent;  coloration  variable,  either  uniform, 

banded,  or  spotted,  or  somewhat  darker 

KEY  TO  THE  OPHICHTHID  EELS  OF  dorsally                                                        9 

JOHNSTON  AND  THE  HAWAIIAN  ISLANDS  8a    Posterior  nostril  opening  outside  mouth, 

la.  Caudal  fin  rays  conspicuous,  confluent  with  a  flap;  anterior  nostril  tubular;  body 

with  dorsal  and  anal;  tail  tip  flexible;  gill  extremely  elongate;  head  15-20  times  in 

openings  mid-lateral,  a  constricted  open-  TL Apterichtus  flavicaudus 

ing;  pectoral  fin  absent  in  Hawaiian  8b.  Posterior  nostril  opening  inside  mouth; 

species.  Subfamily  MYROPHINAE  2  anterior  nostril  flush  with  snout;  body 

Ib.  Tail  tip  a  hard  or  fleshy,  finless  point;  gill  moderately  elongate;  head  11-12  times 

openings  mid-lateral  to  entirely  ventral,  in  TL  Ichthyapus  vulturis 

unconstricted;  pectoral  fin  present  in  9a.  Only  fin  a  short  dorsal  originating  just 

some    species.    Subfamily    OPHICHTHI-  behind  occiput  and  ending  in  anterior 

NAE  7  trunk  region;  body  extremely  elongate, 

2a.  A  prominent  median  toothed  groove  on  the  depth  120-150  times  in  TL 

ventral  side  of  snout,  bordered  by  der-  Phaenomonas  cooperae 

mal  folds,  extending  forward  to  anterior  9b.  Dorsal  and  anal  fins  present,  the  dorsal 

nostrils;  anterior  nostrils  elongated  tubes  extending  nearly  to  the  tail  tip;  body 

equal  to  eye  in  length moderately  to  extremely  elongate,  but 

Schismorhynchus  labialis  the  depth  less  than  120  times  in  TL  __     10 

2b.  Ventral  side  of  snout  without  a  promi-  lOa.  Dorsal  fin  orgin  (DFO)  on  nape;  pectoral 
nent  groove  bordered  by  dermal  folds;  fins  absent;  gill  openings  inferior,  con- 
anterior  nostrils  less  than  eye  in  length  verging  forward;  coloration  pale  to  tan, 

3  overlain  with  small  dark  spots  

3a.  Teeth   absent  on   vomer,   absent   or  Callechelys  luteus 

embedded  on  intermaxillary,  those  on  lOb.  DFO  behind  nape,  either  on  head  or 

maxillary  and  dentary  minute  or  villi-  slightly  behind  gill  openings;  pectoral 

form;  dorsal  fin  origin  (DFO)  behind  anus  fins  present;  coloration  either  uniform, 

Schultzidia  johnstonensis  spotted,  or  banded   11 

3b.  Teeth  present  on  intermaxillary,  maxil-  lla.  DFO  well  in  advance  of  gill  openings; 

lary,  dentary,  and  vomer;  DFO  either  teeth  molariform  or  granular;  pectoral 

before  or  behind  anus  4  fins  broad-based,  short  and  rounded  __  12 

4a.  Posterior  nostril  entirely  outside  of  lib.  DFO  before,  above,  or  behind  gill  open- 
mouth;  teeth  on  maxillary,  dentary,  and  ings;  teeth  pointed;  pectoral  fin  base  re- 


McCOSKER:  HAWAIIAN  SNAKE  EELS 


59 


stricted,  opposite  upper  half  of  gill  open-  ed  by  pale  interspaces;  vertebrae  of  1 

ings  and  longer  than  broad  13  specimen  148 

12a.  Coloration  consists  of  several  longitudi-  Ophichthus  polyophthalmus 

nal  series  of  dark  spots  along  sides  and  19b.  Head  and  body  overlain  with  numerous 

dorsal  surface Myrichthys  maculosus  dark  spots;  those  on  body  in  2  irregular 

12b.  Coloration  consists  of  about  30  dark  sad-  rows,  the  spots  about  equal  in  size  to 

dies  reaching  approximately  to  the  lat-  their  interspaces;  vertebrae  of  6  speci- 

eral  line Myrichthys  bleekeri  mens  152-155 Ophichthus  erabo 

13a.  DFO  well  ahead  of  gill  openings;  edge  of 

upper  lip  fringed  with  a  conspicuous  row  Muraenichthys  puhioilo,  new  species 

of  barbels   Cirrhimurae na  playfairii  (Figures  1-2) 

13b.   DFO  above  or  behind  gill  openings;  up-  Hobtype:  CAS  29115  (originally  SIO  70-32),  355  mm  total 

per  lip  either  naked  or  fringed 14  length,  captured  in  a  benthic  shrimp  trap  set  overnight  at  275 

14a.  Postorbital  region  with  a  conspicuous  m  depth-  N  of  Barber's  Point'  Oahu-  Hawaiian  islands,  by 

..        ...  Thomas  A.  Clarke  on  28  October  1969. 

transverse  depression;  lips  fringed;  ca- 
nine teeth  in  jaws  and  on  vomer;  color-  COUNTS  AND  MEASUREMENTS  (in  mm). — The 
ation  uniform 15  description  of  this  new  species  is  based  on  the 

14b.  Dorsolateral  profile  on  head  even;  lips  holotype  and  only  known  specimen.  Total  length 

entire;  jaw  and  vomerine  teeth  not  ex-  355  mm;  head  length  37.3;  trunk  length  120;  tail 

cessively  developed;  coloration  uniform,  length  198;  body  depth  at  gill  openings  15;  body 

spotted,  or  banded  16  width  at  gill  openings  8;  body  depth  at  anus  12; 

15a.  Dorsal  fine  pale;  snout  contained  about  body  width  at  anus  7.5;  origin  of  dorsal  fin  86; 

15  times  in  head  length snout  length  8.2;  upper  jaw  length  10.7;  eye  di- 

Brachysomophis  sauropsis  ameter  3.2;  interorbital  distance  4.4.  Total  ver- 

15b.  Dorsal  fin  dark  with  a  white  border;  tebrae  160;  preanal  vertebrae  62. 

snout  contained  about  10  times  in  head  GENERAL   DESCRIPTION. — Body   elongate, 

length Brachysomophis  henshawi  depth  at  gill  openings  23.7  in  total  length,  taper- 

16a.  Conspicuous  leaflike  appendages  on  an-  ing  and  laterally  compressed  posteriorly.  Head 

terior  nostrils;  head  and  trunk  equal  to  and  trunk  2.26  and  head  9.5  in  total  length, 

or  greater  than  tail  (coloration  uniform;  Snout  moderately  acute;  lower  jaw  included,  its 

vomerine  teeth  absent)  tip  reaching  the  posterior  edge  of  anterior  nos- 

Phyllophichthus  xenodontus  trils.  Anterior  nostrils  tubular,  slightly  shorter 

16b.  No  leaflike  appendages  on  anterior  nos-  than  eye  diameter.  Posterior  nostril  entirely  in- 

trils;  head  and  trunk  equal  to  or  less  than  side  upper  lip,  before  anterior  margin  of  the  eye, 

tail 17  opening  inward,  appearing  externally  as  a  flap. 

17a.  Vomerine  teeth  absent  or  1-3;  head  and  Eye  less  than  twice  in  fleshy  interorbital  dis- 

body  coloration  light  to  tan,  overlain  tance.   Interorbital  region  flat.   Rictus  of  jaw 

with  a  series  of  black  saddles slightly  behind  posterior  margin  of  eye. 

Leiuranus  semicinctus  Median  fins  low,  except  near  tail  tip  and  an- 

17b.  A  series  of  teeth  on  the  vomer;  color-  terior  portion  of  anal  fin.  Dorsal  fin  arises  closer 
ation  uniform  or  spotted,  not  as  above  to  gill  openings  than  to  anus.  Paired  fins  absent. 
18  Median  fins  confluent  with  caudal. 

18a.  DFO  above  pectoral  tips;  pectoral  fin  Head  pores  minute.  Single  temporal  and  in- 
elongate,  attenuate;  coloration  uniform,  terorbital  pores  present.  Five  pores  along  man- 
darker  dorsally  dible,  three  overlying  preopercle.  Lateral-line 

Ophichthus  kunaloa  n.sp.  pores  difficult  to  discern  posteriorly;  10  pores 

18b.  DFO  above  gill  openings;  pectoral  fin  before  gill  opening;  approximately   140  pores 

rounded;  coloration  not  uniform,  mark-  along  left  side,  65  before  the  anus.  Last  lateral- 

edly  spotted 19  line  pore  occurs  about  a  head  length  before  tail 

19a.  Head  and  body  overlain  with  numerous  tip. 

ocellated  spots;  those  on  body  in  3  reg-  Teeth  slender,  small  and  conical,  uniserial 

ular  alternating  rows,  the  spots  separat-  throughout.  The  pattern  of  dentition  is  nearly 


60 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  2 


FIGURES  1  and  2.    Fig.  1.  Left  lateral  view  of  holotype  of  Muraenichthys  puhioilo  McCosker,  new  species,  CAS  291 15,  355 
mm  TL.  Fig.  2.  Head  region  of  holotype  of  Muraenichthys  puhioilo  McCosker,  new  species. 


identical  to  that  of  Muraenichthys  chilensis  (cf. 
McCosker  1970,  fig.  4),  except  that  the  new 
species  lacks  the  anteriormost  intermaxillary 
tooth.  None  is  extremely  elongate  although  the 
anteriormost  vomerine  tooth  is  slightly  larger 
than  all  jaw  teeth.  Four  intermaxillary  teeth, 
forming  an  inverted  v,  are  followed  by  14  uni- 
serial  vomerine  teeth.  The  maxillary  teeth  are 
equal  in  size,  15  right  and  17  left.  Teeth  of  lower 
jaw  separated  at  symphysis,  about  17  on  each 
side. 

Body  coloration  in  isopropyl  alcohol  uniform 
tan,  although  the  belly  and  lateral-line  are  slight- 
ly darker.  Median  fins  pale  except  for  the  pos- 
terior portion  (slightly  longer  than  head  length) 


of  anal  fin  which  is  dark.  (The  functional  signif- 
icance of  this  highly  contrasting  fin  coloration  is 
not  known.)  Eyes  dark  blue. 

ETYMOLOGY. — From  the  Hawaiian  puhi  oilo, 
small  eels  about  as  large  in  diameter  as  a  finger, 
here  considered  a  noun  in  apposition.  Eels,  par- 
ticularly puhi  oilo,  were  highly  esteemed  as  food 
by  ancient  Hawaiians.  Mary  Kawena  Pukui 
(1902)  wrote  that  "the  eel  was  a  fish  of  which 
chiefs  were  fond  ...  so  much  prized  by  those 
of  Koolau,  Maui  .  .  .  that  they  said  only  beloved 
guests  were  served  with  eels  ...  for  eels  were 
considered  choicer  than  wives." 

REMARKS. — This  individual  was  captured  at 
a  depth  far  greater  than  that  of  any  previously 


McCOSKER:  HAWAIIAN  SNAKE  EELS 


61 


known  Muraenichthys.  The  nearly  20  species  of 
the  genus  are  generally  known  from  water  shal- 
lower than  50  meters. 

This  specimen  was  reported  by  Clarke 
(1972:312),  on  the  basis  of  my  erroneous  iden- 
tification, as  Muraenichthys  macropterus 
Bleeker.  I  have  subsequently  examined  a  radio- 
graph of  Bleeker's  type-specimen  (BM 
1867.11.28.303)  and  found  it  to  possess  130  ver- 
tebrae, with  22  before  the  dorsal  fin  origin  and 
47  before  the  anal  fin  origin.  Bleeker's  type  was 
from  Ambon;  a  series  from  Palau  (CAS  41186) 
had  127-132  vertebrae  (jc  =  129.8,  n  =  5).  My 
examination  of  more  than  100  specimens  of  M. 
macropterus  from  throughout  Oceania  found 
them  to  differ  from  the  new  species  in  having 
fewer  vertebrae,  biserial  vomerine  dentition  (be- 
coming uniserial  posteriorly),  uniform  fin  col- 
oration, and  in  occupying  shallower  water.  The 
new  species  differs  from  all  other  species  of 
Muraenichthys,  subgenus  Scolecenchelys,  on 
the  basis  of  its  uniserial  dentition,  anterior  dor- 
sal fin  location,  coloration,  and  vertebral  num- 
ber. 

In  my  review  of  Muraenichthys  (McCosker 
1970),  I  followed  Schultz  (1953)  in  considering 
M.  breviceps  Giinther  to  be  a  probable  synonym 
of  M.  macropterus.  I  have  subsequently  rec- 
ognized M.  breviceps  as  a  distinct  species  and 
include  M.  devisi  Fowler,  M.  ogilbyi  Fowler, 
and  Aotea  acus  Phillipps  in  its  synonymy 
(McCosker  and  Allen,  MS.).  I  also  examined  the 
other  known  synonyms  of  M.  macropterus,  M. 
owstoni  Jordan  and  Snyder  from  Japan  and 
Echidna  uniformis  Scale  from  Guam,  and  de- 
termined that  they  are  M.  macropterus.  The  ho- 
lotype  of  M.  owstoni  (SU  6472)  has  131  verte- 
brae. 

Ophichthus  kunaloa,  new  species 

(Figures  3-4) 

Holotype:  CAS  29136  (originally  SIO  70-33),  440  mm  total 
length,  captured  in  a  benthic  shrimp  trap  set  overnight  at 
350-m  depth,  SE  of  Barber's  Point,  Oahu,  Hawaiian  Islands, 
by  Thomas  A.  Clarke  on  31  December  1969. 

Paratypes:  Collected  with  the  holotype.  BPBM  21086,  473 
mm  total  length.  USNM  218274,  only  the  head  and  anterior 
trunk  region  of  a  partially  eaten  specimen. 

COUNTS  AND  MEASUREMENTS  (in  mm). — The 
condition  of  the  holotype  is  parenthetically  fol- 
lowed by  that  of  the  intact  paratype.  Total  length 
440  mm  (473  mm);  head  length  42.0  (45.7);  trunk 
length  135  (144);  tail  length  263  (283);  body  depth 


at  gill  openings  16  (15);  body  width  at  gill  open- 
ings 13.7  (12.5);  origin  of  dorsal  fin  58  (68);  left 
pectoral  fin  length  20.0  (19.5);  left  pectoral  fin 
base  4.6  (4.5);  snout  length  8.6  (7.7);  upper  jaw 
length  17.2  (20.0);  gill  opening  height  5.6  (5.5); 
eye  diameter  7.0  (8.0);  interorbital  distance  6.9 
(6.9).  Total  vertebrae  185  (181);  preanal  verte- 
brae 66  (67). 

GENERAL  DESCRIPTION. — Body  elongate, 
depth  at  gill  openings  10.3-10.5  in  total  length, 
tapering  posteriorly.  Tail  slender,  laterally  com- 
pressed posteriorly.  Head  and  trunk  2.49  and 
head  10.3-10.5  in  total  length.  Snout  rounded; 
lower  jaw  included,  its  tip  in  advance  of  anterior 
nostril  base.  Anterior  nostril  tubular;  posterior 
nostril  at  edge  of  lip,  entirely  outside  of  mouth, 
covered  anteriorly  by  a  small  flap.  Eye  large;  its 
center  lies  slightly  behind  midpoint  of  upper  jaw. 
Head  broad;  interorbital  area  flat. 

Median  fins  low,  lying  partially  within  a 
groove.  Dorsal  fin  arises  before  end  of  pectoral. 
Median  fins  disappear  within  a  flabby  groove 
before  the  tail  tip.  Pectoral  fins  elongate,  the 
dorsalmost  rays  tapering  posteriorly.  Caudal  tip 
naked. 

Head  pores  minute,  difficult  to  locate  on  pre- 
served specimens.  Single  temporal  and  interor- 
bital pores.  Five  pores  along  mandible,  two 
overlying  preopercle.  Lateral-line  pores  be- 
gin above  second  preopercular  pore.  Lateral- 
line  pores  difficult  to  discern,  about  64  before 
anal  opening. 

Teeth  small,  conical;  biserial  in  jaws,  the  out- 
er row  smaller  and  closer  set.  Vomerine  teeth 
biserial  anteriorly,  followed  by  a  uniserial  row 
of  about  10  teeth.  Intermaxillary  tooth  patch 
surrounded  anterolaterally  by  a  ring  of  6-8 
teeth. 

Body  coloration  in  isopropyl  alcohol  tan,  be- 
coming lighter  along  chin,  snout,  throat,  lower 
third  of  trunk  region,  below  the  lateral-line  of 
the  trunk  region,  along  the  dorsal  midline,  and 
at  the  tail  tip.  A  black  smudge  exists  along  the 
lower  edge  of  the  tail,  about  l/s  head  length  from 
the  tail  tip.  Fins  pale.  Lateral-line  pores  lie  with- 
in minute  white  dots.  Eyes  blue. 

ETYMOLOGY. — Named  kunaloa  in  reference 
to  Kuna  Loa,  the  Long  Eel,  of  ancient  Hawaiian 
legends.  It  is  said  that  the  sixth  great  deed  of 
Maui  the  Wonder  Boy  was  to  behead  Kuna  Loa 
after  the  treacherous  eel  had  assaulted  the  fair 
maiden  Hina  (see  Colum  1937).  The  legend  as- 


62 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  2 


FIGURES  3  and  4.     Fig.  3.  Left  lateral  view  of  holotype  of  Ophichthus  kunaloa  McCosker,  new  species,  CAS  29136,  440  mm 
TL.  Fig.  4.  Head  region  of  holotype  of  Ophichthus  kunaloa  McCosker,  new  species. 


serts  that  from  the  cut  tail  evolved  the  common 
conger  eel  and  from  the  blood  which  fell  into  the 
fresh  and  salt  water  came  all  of  the  other  Hawai- 
ian eels.  This,  clearly,  was  the  first  attempt  at 
a  phylogenetic  interpretation  of  Hawaiian  an- 
guilliforms. 

REMARKS. — The  depth  of  capture  of  the  new 
species  is  remarkable  in  that  most  benthic 
ophichthids  live  shallower  than  100  meters.  It  is 
entirely  likely  that  future  deepwater  collections 
in  outlying  areas  might  reveal  this  or  a  closely 
related  species. 

In  his  key  to  the  Hawaiian  ophichthids,  Gos- 
line  (1951:309)  mentioned  a  new  species  of  oph- 
ichthine  which  possessed  characters  similar  to 
the  new  species,  that  was  "probably  from  mod- 
erately deep  water,"  and  had  been  killed  by  the 
Mauna  Loa  lava  flow  of  1950.  This  specimen 
was  not  mentioned  in  further  publications,  and 
neither  Gosline  (in  litt.),  John  E.  Randall  of  the 
Bishop  Museum  (BPBM),  nor  Leighton  Taylor 
of  the  University  of  Hawaii  (UH)  have  been  able 
to  locate  it  after  the  majority  of  the  UH  fish 
collection  had  been  transferred  to  the  BPBM. 

The  closest  relatives  to  the  new  species  appear 
to  be  those  congeners  which  also  possess  large 
eyes,  similar  dentition,  posterior  nostrils  along 
the  lip  (rather  than  opening  into  the  mouth)  and 


preceded  by  a  flap,  two  rather  than  three  pre- 
opercular  pores,  and  a  plain  coloration.  Those 
species  of  Ophichthus,  mostly  within  the  sub- 
genus  Coecilophis  Kaup  (cf.  McCosker  1977), 
share  a  preference  for  moderate-depth  sand  or 
mud  substrates.  Ophichthus  kunaloa  is  partic- 
ularly similar  to  O.  urolophus  (Temminck  and 
Schlegel),  an  oriental  species,  which  differs  in 
its  proportionately  shorter  tail  and  much  deeper 
body,  and  to  the  eastern  Pacific  O.  pacific! 
Giinther,  a  species  with  a  comparatively  longer 
head,  deeper  body,  and  white  spots  along  the 
lateral-line. 

The  following  comments  comprise  new  rec- 
ords and  systematic  information  concerning  the 
Hawaiian  ophichthid  fauna. 

SUBFAMILY  MYROPHINAE 
Sdiismorhv iichus  lahialis  (Scale) 

Muraenichthys  labialis   Scale,    1917:79  (type-locality,   Arno 

Atoll,  Marshall  Islands). 
Leptenchelys  labialis:  Schultz  1953:80. 
Schismorhynchus  labialis:  McCosker  1970:509. 

REMARKS. — This  wide-ranging  species  has 
been  collected  from  the  Society  Islands  (BPBM 
12016),  Marshall  Islands,  Johnston  Island,  and 
Easter  Island,  but  has  not  been  captured  in  Ha- 
waii. 


McCOSKER:  HAWAIIAN  SNAKE  EELS 


63 


Muraenichthys  cookei  Fowler 

Muraenichthys  cookei  Fowler,  1928:41  (type-locality,  Oahu). 

REMARKS. — I  concur  with  Gosline  (1951),  that 
M.  cookei,  a  Hawaiian  endemic,  is  closely  re- 
lated to  but  distinct  from  M.  laticaudata.  A 
comprehensive  discussion  of  specific  differences 
exists  in  Gosline  (1955:469^70). 

SUBFAMILY  OPHICHTHINAE 
Apterichtus  flavicaudus  (Snyder) 

Sphagebranchus  flavicaudus  Snyder,  1904:516  (type-locality, 

between  Maui  and  Lanai). 

Caecula  (Sphagebranchus)  flavicauda:  Gosline  1951:311. 
Verma  flavicauda:  Bohlke  1968:3. 
Apterichtus  flavicaudus:  Bohlke  and  McCosker  1975:4. 

REMARKS. — This  species,  previously  consid- 
ered a  Hawaiian  endemic,  has  now  been  col- 
lected at  several  South  Pacific  locations.  I  com- 
pared all  of  Snyder's  specimens  with  specimens 
from  Midway  Island  (SIO  68^87)  and  Rapa  Is- 
land (BPBM  12306),  and  found  them  to  differ 
only  in  vertebral  number.  Six  Hawaiian  and 
Midway  specimens  had  155-166  (x  =  158.7)  ver- 
tebrae, whereas  six  Rapa  specimens  had  163- 
166  (jf  -  164). 

Ichthyapus  vulturis  (Weber  and  de  Beaufort) 

Sphagebranchus  vulturis  Weber  and  de  Beaufort,   1916:319 

(type-locality,  Nasi  besar  Island,  Sumatra). 
Caecula  (Sphagebranchus)  platyrhyncha  Gosline,    1951:312 

(type-locality,  Oahu,  Hawaiian  Islands). 

REMARKS. — Randall  and  McCosker  (1975) 
synonymized  Caecula  platyrhyncha  with 
Sphagebranchus  vulturis  after  comparing 
Hawaiian  specimens  with  the  holotype.  Varia- 
tion exists  in  the  number  of  preopercular  pores 
(either  3  or  4)  of  Hawaiian  specimens,  but  the 
pore  number  of  specimens  from  other  localities 
was  found  to  be  constant. 

Phaenomonas  cooperae  Palmer 

Phaenomonas  cooperae  Palmer,  1970:219  (type-locality,  Gil- 
bert Islands). 

REMARKS. — This  unmistakable  elongate 
species  is  known  from  Hawaii  on  the  basis  of  a 
specimen  (HIMB  68-52)  dredged  from  a  depth 
of  60  m,  offshore  from  Keehi  Lagoon,  Oahu,  in 
1968.  The  specimen  has  262  total  vertebrae,  169 
preanal,  and  falls  within  the  range  of  meristic 
and  morphometric  variation  of  its  Indo- West- 
Pacific  conspecifics  (McCosker  1975). 


Callechelys  luteus  Snyder 

Callechelys  luteus  Snyder,  1904:517  (type-locality,  near  the 
southern  coast  of  Molokai). 

REMARKS. — This  elongate  species,  unique  in 
dorsal  fin  condition  and  coloration,  is  known 
only  from  the  Hawaiian  Islands  and  Midway  Is- 
land (SIO  68-497).  Its  closest  relative  appears 
to  be  the  eastern  Pacific  endemic  C.  galapagen- 
sis  McCosker  and  Rosenblatt.  A  radiograph  of 
the  holotype  of  C.  luteus  (USNM  50864)  evi- 
denced 216  vertebrae,  with  123  before  the  anal 
fin  origin. 

Myrichthys  maculosus  (Cuvier) 

Muraena  maculosa  Cuvier,  1817:232  (type-locality,  European 
Seas?). 

Pisoodonophis  magnifica  Abbott,  1861:476  (type-locality, 
Hawaiian  Islands). 

Ophichthus  stypurus  Smith  and  Swain,  1882:120  (type-locali- 
ty, Johnston  Island). 

REMARKS. — This  common  species,  perhaps 
better  than  any  other  Hawaiian  ophichthid,  de- 
picts the  isolation  of  the  Hawaiian  and  Johnston 
population  as  evidenced  by  vertebral  number. 
I  am  unable  except  by  vertebral  number  to  sep- 
arate the  Hawaiian,  Midway,  and  Johnston 
specimens  from  those  from  the  Red  Sea,  Indian 
Ocean,  Oceania,  and  the  eastern  tropical  Pacific 
(see  Table  1).  For  this  and  related  faunal  studies 
I  have  examined  many  living  and  dead  speci- 
mens of  Myrichthys  as  well  as  the  types  of  most 
of  the  nominal  species.  Until  now,  I  have  fol- 
lowed the  conventional  view  that  the  eastern 
Pacific  nominal  species  M.  xysturus  (Jordan  and 
Gilbert)  (which  includes  M.  tigrinus  Girard  and 
M.  pantostigmius  Jordan  and  McGregor)  differs 
from  M.  maculosus,  although  the  only  signifi- 
cant difference  is  that  of  vertebral  number.  In 
the  absence  of  any  apparent  morphological  dif- 
ferences, I  am  resigned  to  recognize  these  pop- 
ulations at  a  subspecific  level,  and  therefore  rec- 
ognize this  ophichthid  as  a  trans-Pacific  species. 
The  differences  in  M.  maculosus  population 
vertebral  numbers  is  thus  parallel  to  the  condi- 
tion of  several  trans-Pacific  species  of  muraenid 
eels  (Rosenblatt  et  al.  1972;  McCosker  and  Ro- 
senblatt 1975).  I  suspect  that  as  data  are  gath- 
ered concerning  the  duration  of  larval  life,  dis- 
tance of  larval  transport,  and  the  critical  period 
of  leptocephalus  transformation,  a  more  lucid 
view  of  Myrichthys  taxonomy  will  be  achieved. 

I  have  examined  the  holotypes  of  M.  magni- 


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TABLE  1.  Myrichthys  maculosus  (CuviER):  VERTEBRAL 
DATA  FOR  EASTERN  PACIFIC  AND  INDO-PACIFIC  POPULA- 
TIONS. All  counts  made  from  radiographs  of  adults. 


mean 

range 

n 

Eastern  Pacific1 

153.1 

149-157 

19 

Revillagigedos  and 

Clipperton  Is. 

163.1 

161-168 

10 

Hawaii,  Johnston,  and 

Midway 

178.5 

174-182 

16 

Western  Pacific2 

192.8 

190-199 

8 

Red  Sea 

194.5 

194-195 

2 

1  Specimens  from  the  Gulf  of  California,  Tres  Marias  Is- 
lands, Cocos  Island,  Panama,  and  the  Galapagos  Islands. 

2  Specimens  from   Wake,   Palau,   Philippines,   RyuKyus, 
and  the  Line  Islands. 


ficus  and  M.  stypurus  and  found  them  to  be  con- 
specific.  The  holotype  of  M.  stypurus  is  aberrant 
in  that  its  tail  had  been  severed  and  subsequent- 
ly healed. 

Myrichthys  bleekeri  Gosline 

Ophisurus  fasciatus  var.  semicinctus  Bleeker,  1864:64  [a 
homonym  of  Ophisurus  semicinctus  Lay  and  Bennett, 
1839:66]  (type-locality,  Indonesia). 

Myrichthys  bleekeri  Gosline,  1951:314  [a  substitute  name  for 
Ophisurus  fasciatus  var.  semicinctus  Bleeker,  1864,  preoc- 
cupied]. 

REMARKS. — This  species  is  known  from  John- 
ston Island  and  the  Indo-West-Pacific,  but  not 
from  Hawaii. 

Cirrhimuraena  playfairii  (Giinther) 

Ophichthys  playfairii  Giinther,  1870:76  (type-locality,  Zanzi- 
bar). 

Microdonophis  macgregori  Jenkins,  1903:422  (type-locality, 
Lahaina,  Maui). 

Jenkinsiella  macgregori:  Jordan  and  Evermann,  1905:82. 

Cirrhimuraena  playfairii:  Barnard,  1925:205. 

Cirrhimuraena  macgregori:  Gosline,  1951:315. 

REMARKS. — Smith  (1962)  synonymized  the 
Hawaiian  species  Microdonophis  macgregori 
with  the  wide-ranging  Indo-Pacific  species  Cir- 
rhimuraena playfairii.  He  based  this  action  on 
his  comparison  of  four  specimens  from  Aldabra 
with  published  descriptions  by  Gosline  (1951)  of 
Hawaiian  specimens  and  by  Weed  and  Howarth 
(1961)  of  specimens  from  Ceylon.  I  have  ex- 
amined the  type-specimens  from  Hawaii  and 
compared  them  with  material  from  Midway  (SIO 
68^97)  and  the  Marquesas  (BPBM  12304),  and 
concur  with  Smith.  The  holotype  of  M.  mac- 
gregori (USNM  50721)  has  180  vertebrae;  two 


Marquesan  specimens  have  186  and  187  verte- 
brae. 

Brachysomophis  sauropsis  Schultz 

Brachysomophis  sauropsis  Schultz,   1943:18  (type-locality, 
Samoa). 

REMARKS. — Not  known  from  Hawaii.  I  have 
been  unable  to  examine  Gosline's  (1955:443) 
specimen  of  B.  sauropsis  from  Johnston  Island. 
However,  based  on  his  description,  I  would 
agree  that  B.  sauropsis  and  B.  henshawi  are  dis- 
tinct species. 

Brachysomophis  henshawi  Jordan  and  Snyder 

Brachysomophis  henshawi  Jordan  and   Snyder,    1904:940 
(type-locality,  Honolulu). 

REMARKS. — Apparently,  this  species  is  a 
Hawaiian  endemic.  A  comprehensive  revision 
of  Brachysomophis  is  necessary,  however,  be- 
fore assumptions  concerning  its  species  can  be 
made. 

Phyllophichthus  xenodontus  Gosline 

Phyllophichthus  xenodontus  Gosline,  1951:316  (type-locality, 
Oahu). 

REMARKS. — Phyllophichthus  is  currently  rec- 
ognized to  contain  a  single  wide-ranging  species, 
known  from  the  western  Indian  Ocean,  through- 
out Oceania,  and  Hawaii  (McCosker  and  Allen, 
MS).  Four  specimens  from  Johnston  and  Oahu 
islands  had  168-175  vertebrae  (Jc  =  170.8). 

Ophichthus  polyophthalmus  Bleeker 

(Figure  5) 

Ophichthys  polyophthalmus  Bleeker,   1864:43  (type-locality, 

Ambon). 

Microdonophis  polyophthalmus:  Jordan  and  Snyder  1907:207. 
Ophichthus  polyophthalmus:  Fowler  1927:5. 

REMARKS. — The  markedly  spotted  Hawaiian 
eels  placed  by  recent  authors  in  either  Micro- 
donophis or  Ophichthus  is  a  composite.  Oph- 
ichthus polyophthalmus  (Fig.  5),  a  medium- 
length  adult  ophichthid  with  ocellated  spots  on 
the  head  and  body,  was  reported  by  Fowler 
(1927)  on  the  basis  of  a  Kahoolawe  (BPBM  3700) 
dredge  specimen.  Fowler  (1928)  subsequently 
recorded  eleven  Honolulu  market  specimens 
(the  largest,  1,039  mm)  which  possessed  dark 
spots;  he  considered  them  to  also  be  O.  poly- 
ophthalmus and  recognized  the  nominal  species 
Microdonophis  fowled  Jordan  and  Evermann 


McCOSKER:  HAWAIIAN  SNAKE  EELS 


FIGURE  5.     Adult  specimen  of  Ophichthus  polyophthalmus,  taken  from  Bleeker  (1864). 


and  Ophichthus  garretti  Giinther  to  be  syn- 
onyms which  differed  only  in  coloration.  Oph- 
ichthus garretti  is  a  valid  and  distinctly  different 
species.  My  comparison  of  numerous  specimens 
of  the  large  form  with  solid  dark  spots  (Ophich- 
thus erabo,  Fig.  6)  with  the  medium-length  eel 
with  ocellated  spots  indicated  that  they  are  in 
fact  separate  species  differing  in  coloration  and 
vertebral  number.  Too  few  specimens  of  O. 
polyophthalmus  were  available  for  a  proper 
morphological  comparison,  although  O.  erabo 
appears  to  possess  a  proportionately  longer  tail. 
I  herein  report  a  second  Hawaiian  specimen 
of  O.  polyophthalmus,  collected  in  1968  by  hook 
and  line  from  Nanakuli,  Oahu  (BPBM  1 1981,  399 
mm  SL).  The  specimen  has  148  vertebrae,  75 
before  the  anal  opening. 

Ophichthus  erabo  (Jordan  and  Snyder) 

(Figure  6) 

Microdonophis  erabo  Jordan  and  Snyder,  1901:780  (type-lo- 
cality, Misaki,  Japan). 

Microdonophis  fowleri  Jordan  and  Evermann,  1904:164  (type- 
locality,  Honolulu). 

Ophichthus  erabo:  McCosker  1977:81. 

REMARKS. — The  majority  of  Hawaiian  rec- 
ords of  O.  polyophthalmus  are  based  on  speci- 
mens of  O.  erabo.  I  have  compared  the  Japa- 
nese holotype  (SU  6477)  and  cotypes  (SU  6667 


and  6744)  of  O.  erabo  with  specimens  from  Ha- 
waii (SU  8407;  SU  8466;  BPBM  12510;  and 
USNM  50613,  the  holotype  of  M.  fowleri)  and 
Taiwan  (CAS  15600),  and  found  them  not  to  dif- 
fer. The  vertebral  range  of  six  specimens  was 
152-155  (x  =  154).  The  holotype  of  O.  erabo  has 
155  vertebrae  and  the  holotype  of  M.  fowleri  has 
152. 

DISCUSSION 

The  endemism  of  the  Hawaiian  shorefish  fau- 
na has  been  recognized  by  numerous  authors 
and  most  recently  summarized  by  Randall 
(1976).  He  calculated  that  29  percent  of  the  442 
reef  and  shorefish  species  were  endemic  at  the 
specific  or  subspecific  level.  He  elucidated  the 
situation  of  the  problematical  species  such  as 
Acanthurus  triostegus,  wherein  an  identifiable 
difference  in  coloration  exists  between  the 
Hawaiian  and  extralimital  populations,  but  a 
consensus  of  opinion  concerning  the  biological 
significance  of  that  difference  has  not  been 
reached.  A  similar  problem  exists  with  the 
Hawaiian  ophichthids. 

Of  the  15  ophichthids  present  at  Hawaii,  five 
species  are  endemic.  The  new  species  described 
herein,  Ophichthus  kunaloa  and  Muraenichthys 
puhioilo,  are  known  only  from  a  single  deep- 
water  collection  and  are  therefore  of  little  value 


FIGURE  6.     Adult  specimen  of  Ophichthus  erabo,  taken  from  Jordan  and  Snyder  (1901). 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  2 


TABLE  2.   DISTRIBUTION  OF  HAWAIIAN  AND  JOHNSTON  ISLAND  OPHICHTHIDS. 


Hawaiian 


Leewards 


Johnston 


Indo-West 
Pacific 


Sihultzidia  johnstonensis 
Schismorhynchus  labialis 
Muraenichthys  schultzei 
Muraenichthys  cookei 
Muraenichthys  gymnotus 
Muraenichthys  puhioilo 
Ichthyapus  vulturis 
Apterichtus  flavicaudus 
Callechelys  luteus 
Myrichthys  maculosus 
Myrichthys  bleekeri 
Cirrhimuraena  playfairii 
Phyllophichthus  xenodontus 
Phaenomonas  cooperae 
Leiuranus  semicinctus 
Brachysomophis  sauropsis 
Brachysomophis  henshawi 
Ophichthus  erabo 
Ophichthus  polyophthalmus 
Ophichthus  kunaloa 


to  a  zoogeographic  analysis.  The  endemics,  Mu- 
raenichthys cookei  and  Brachysomophis  hen- 
shawi, are  perhaps  no  more  different  than  are 
several  of  the  other  ophichthids  discussed  here 
from  their  Indo-West-Pacific  "conspecifics." 
Callechelys  luteus  is  the  only  Hawaiian 
ophichthid  endemic  distinctly  different  at  the 
specific  level  from  all  known  congeners.  It  is 
most  closely  related  to  C.  galapagensis,  another 
insular  endemic  (McCosker  and  Rosenblatt 
1972).  In  my  analysis  of  Hawaiian  ophichthids, 
I  have  been  able  to  recognize  the  Hawaiian  pop- 
ulations of  several  species  (viz.,  Myrichthys 
maculosus,  Phaenomonas  cooperae,  Phylloph- 
ichthus xenodontus,  Ichthyapus  vulturis,  and 
probably  several  others)  to  be  distinct  from  oth- 
er Indo-Pacific  populations  of  their  conspecifics 
on  the  basis  of  vertebral  differences.  Yet  I  am 
hesitant  to  assign  them  specific  rank.  It  should 
be  recognized,  however,  that  the  absolute  dif- 
ferences in  ophichthid  vertebral  numbers  appear 
to  be  less  when  calculated  on  a  percentage  basis. 
For  example,  the  difference  of  eight  vertebrae 
between  the  Hawaiian  specimen  of  Phaenomon- 
as cooperae  and  the  Gilbert  Island  holotype  is 
only  three  percent,  which  is  less  than  a  differ- 
ence of  one  vertebra  for  most  perciform  fishes. 
As  Randall  (1976:49-50)  has  discussed,  these 
differences  are  real  and  apparently  indicate  lim- 
ited gene  flow  with  other  insular  populations. 
Whether  the  Hawaiian  forms  are  in  fact  distinct 


biological  species,  incipient  species,  or  what- 
ever taxon  a  systematist  deems  them,  awaits  the 
discovery  of  biological  data  concerning  lepto- 
cephalus  transport,  longevity,  and  gene  flow. 

Untaxing  the  taxonomy  of  the  Hawaiian 
ophichthids,  initiated  by  Maui  the  Wonder  Boy 
and  continued  by  Jordan,  Evermann,  and  Gos- 
line,  remains  a  challenge. 

ACKNOWLEDGMENTS 

This  study  depended  primarily  upon  recent 
specimens  collected  by  J.  E.  Randall  (BPBM), 
the  George  Vanderbilt  Foundation  (CAS),  and 
the  1968  STYX  Expedition  of  the  Scripps  Insti- 
tution of  Oceanography.  Other  specimens  and 
assistance  were  provided  by  the  following  indi- 
viduals: J.  Paxton  and  D.  Hoese  (AMS);  J. 
Bohlke  (ANSP);  A.  Wheeler  (BMNH);  W.  Esch- 
meyer  (CAS);  R.  Lavenberg  (LACM);  R.  Ro- 
senblatt and  J.  Copp  (SIO);  V.  Springer  and  N. 
Gramblin  (USNM);  and  L.  Taylor  of  the  Wai- 
kiki  Aquarium,  Hawaii.  J.  Gordon  and  C.  Ruark 
assisted  with  the  preparation  of  radiographs,  M. 
Giles  prepared  the  photographs,  and  K.  Smith 
prepared  Figures  1-4.  I  am  very  grateful  to  all 
of  the  above-mentioned  individuals  for  their  as- 
sistance, and  in  particular,  to  Thomas  A.  Clarke 
for  providing  the  new  species  and  to  William  A. 
Gosline  for  his  pioneering  studies  of  apodal  fish- 
es. 


McCOSKER:  HAWAIIAN  SNAKE  EELS 


67 


LITERATURE  CITED 

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fishes  in  the  museum  of  the  Academy  of  Natural  Sciences 

of  Philadelphia.  Proc.  Acad.  Nat.  Sci.  Phila.  12:475^*79. 
BARNARD,  K.  H.   1925.  A  monograph  of  the  marine  fishes  of 

South  Africa.  Part  I.  Ann.  S.  Afr.  Mus.  21:1-418. 
BLEEKER,  P.  1864.  Atlas  ichthyologique  des  Indes  Orientales 

Neerlandaises.  Vol.  4.  Amsterdam.  132  pp. 
BOHLKE,  J.  E.    1968.   A  new  species  of  the  ophichthid  eel 

genus  Verma  from  the  West  Atlantic,  with  comments  on 

related  species.  Not.  Nat.  (Phila.),  no.  415.  12  pp. 
,  AND  J.   E.  MCCOSKER.    1975.    The  status  of  the 

ophichthid  eel  genera  Caecula  Vahl  and  Sphagebranchus 

Bloch,  and  the  description  of  a  new  genus  and  species  from 

fresh  waters  in  Brazil.  Proc.  Acad.  Nat.  Sci.  Phila.  127(1): 

1-11. 
CLARKE,  T.  A.  1972.  Collections  and  submarine  observations 

of  deep  benthic  fishes  and  decapod  Crustacea  in  Hawaii. 

Pac.  Sci.  26(3):310-317. 
COLUM,  P.  1937.  Legends  of  Hawaii.  Yale  Univ.  Press,  New 

Haven.  220  pp. 

CUVIER,  G.   1817.  Le  regne  animal.  Vol.  2.  Paris.  532  pp. 
FOWLER,  H.  W.    1927.  Fishes  of  the  tropical  central  Pacific. 

Bernice  P.  Bishop  Mus.,  Bull.  38.  32  pp. 
.  1928.  The  fishes  of  Oceania.  Bernice  P.  Bishop  Mus., 

Mem.  10.  540  pp. 
GOSLINE,  W.  A.  1951.  The  osteology  and  classification  of  the 

ophichthid  eels  of  the  Hawaiian  Islands.  Pac.  Sci.  5(4):298- 

320. 
.    1955.   The  inshore  fish  fauna  of  Johnston  Island,  a 

Central  Pacific  atoll.  Pac.  Sci.  9(4):442-480. 
,  AND  V.  E.  BROCK.  1960.  Handbook  of  Hawaiian  fish- 


genus,  Schismorhynchus,   and  a  new  species,  Muraen- 
ichthys  chilensis.  Pac.  Sci.  24(4):506-516. 
.     1975.    The   eel   genus   Phaenomonas    (Pisces, 


es.  Univ.  Hawaii  Press,  Honolulu.  372  pp. 
GUNTHER,  A.  C.   1870.  Catalogue  of  the  fishes  in  the  British 

Museum.  Vol.  8.  London.  549  pp. 
JENKINS,  O.  P.    1904.  Report  on  collections  of  fishes  made  in 

the  Hawaiian  Islands,  with  descriptions  of  new  species. 

U.S.  Bur.  Fish.  Bull.  22:417-511. 
JORDAN,  D.  S.,  AND  B.  W.  EVERMANN.    1905.  The  aquatic 

resources  of  the  Hawaiian  Islands.  Part  I.  The  shore  fishes. 

U.S.  Fish.  Comm.  Bull.  23(1):  1-574. 
,  AND  J.  O.  SNYDER.    1901.   A  review  of  the  apodal 

fishes  or  eels  of  Japan,  with  descriptions  of  19  new  species. 

Proc.  U.S.  Natl.  Mus.  23(1239):837-890. 
.  1904.  Notes  on  collections  of  fishes  from  Oahu  Island 


and  Laysan  Island,  with  descriptions  of  four  new  species. 
Proc.  U.S.  Natl.  Mus.  27:939-948. 
.    1907.   Notes  on  fishes  of  Hawaii,  with  descriptions 


of  new  species.  U.S.  Bur.  Fish.  Bull.  26:207-218. 

LAY,  G.  T.,  AND  E.  T.  BENNETT.  1839.  The  zoology  of  Cap- 
tain Beechey's  voyage.  Fishes.  London.  Pp.  41-75. 

McCosKER,  J.  E.  1970.  A  review  of  the  eel  genera  Lep- 
tenchelys  and  Muraenichthys,  with  the  description  of  a  new 


Ophichthidae).  Pac.  Sci.  29(4): 36 1-363. 

1977.  The  osteology,  classification,  and  relationships 


of  the  eel  family  Ophichthidae.  Proc.  Calif.  Acad.  Sci.,  Ser. 
4,  41(1):1-123. 
,  AND  R.  H.  ROSENBLATT.    1972.    Eastern  Pacific 


snake-eels  of  the  genus  Callechelys  (Apodes:  Ophichthi- 
dae). Trans.  San  Diego  Soc.  Nat.  Hist.  17(2):  15-24. 
,  AND .    1975.  The  moray  eels  (Pisces:  Muraen- 


idae)  of  the  Galapagos  Islands,  with  new  records  and  syn- 
onymies of  extralimital  species.  Proc.  Calif.  Acad.  Sci.,  Ser. 
4,  40(13):417-427. 

PALMER,  G.  1970.  New  records,  and  one  new  species,  of 
teleost  fishes  from  the  Gilbert  Islands.  Bull.  Brit.  Mus.  (Nat. 
Hist.)Zool.  19(6):2 11-234. 

PUKUI,  M.  K.  1902.  Ka  hana  kuhikuhi  no  ka  lawai'a  ana.  Ka 
Nupepa  Kuokoa,  newspaper  article,  23  May.  (Not  seen; 
reference  from:  TITCOMB,  M.  1972.  Native  use  of  fish  in 
Hawaii.  Univ.  Hawaii  Press,  Honolulu.  175  pp.) 

RANDALL,  J.  E.  1976.  The  endemic  shore  fishes  of  the 
Hawaiian  Islands,  Lord  Howe  Island  and  Easter  Island. 
Colloque  Commerson  1973,  ORSTOM  Trav.  et  Doc.  no. 
47:49-73. 

,  AND  J.  E.  McCosKER.  1975.  The  eels  of  Easter  Island 

with  a  description  of  a  new  moray.  Los  Angeles  County 
Mus.  Contrib.  Sci.,  no.  264.  32  pp. 

ROSENBLATT,  R.  H.,  J.  E.  McCosKER,  AND  I.  RUBINOFF. 
1972.  Indo-West-Pacific  fishes  from  the  Gulf  of  Chiriqui, 
Panama.  Los  Angeles  County  Mus.  Contrib.  Sci.,  no.  234. 
18pp. 

SCHULTZ,  L.  P.  1943.  Fishes  of  the  Phoenix  and  Samoan 
islands.  U.S.  Natl.  Mus.  Bull.  180.  316pp. 

.  1953.  Fishes  of  the  Marshall  and  Marianas  islands. 

Families  from  Asymmetrontidae  through  Siganidae.  U.S. 
Natl.  Mus.  Bull.  202,  1.  685  pp.  (With  collaborators.) 

SEALE,  A.  1917.  New  species  of  apodal  fishes.  Bull.  Mus. 
Comp.  Zool.  Cambridge  61:79-94. 

SMITH,  J.  L.  B.  1962.  Sand-dwelling  eels  of  the  western  In- 
dian Ocean  and  the  Red  Sea.  Rhodes  Univ.  Ichthyol.  Bull. 
24:447-166. 

SMITH,  R.,  AND  J.  SWAIN.  1882.  Notes  on  a  collection  of 
fishes  from  Johnston's  Island  including  descriptions  of  five 
new  species.  Proc.  U.S.  Natl.  Mus.  5:119-143. 

SNYDER,  J.  O.  1904.  A  catalogue  of  the  shore  fishes  collected 
by  the  steamer  "Albatross"  about  the  Hawaiian  Islands  in 
1902.  U.S.  Bur.  Fish.  Bull.  22:513-538. 

WEBER,  M.,  AND  L.  F.  DE  BEAUFORT.  1916.  The  fishes  of 
the  Indo-Australian  Archipelago.  Vol.  3.  Leiden.  455  pp. 

WEED,  W.  A.  Ill,  AND  J.  N.  HOWARTH.  1961.  Range  exten- 
sions of  the  ophichthid  eels  Phyllophichthus  xenodontus 
and  Cirrhimuraena  macgregori.  Copeia  1961(3):357. 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  3,  pp.  69-85;  7  figs. 


December  22,  1979 


REVIEW  OF  THE  RHINOTRAGINI  OF  MEXICO 
(COLEOPTERA:  CERAMBYCIDAE) 

By 

John  A.  Chemsak  and  E.  G.  Linsley 

University  of  California,  Berkeley,  CA  94720 


ABSTRACT:  The  Rhinotragini  known  to  occur  in  Mexico  are  reviewed.  Keys  are  provided  for  the  determination 
of  the  seven  genera  and  also  for  species  in  the  genera.  Distribution  maps  are  included  for  the  more  common 
species  and  most  of  the  new  taxa  are  illustrated.  New  species  include:  Tomopterus  exilis,  Acyphoderes  amoena, 
A.  fulgida,  A.  prolixa,  and  A.  parva. 

New  synonymies  and  one  new  combination  are  proposed. 


INTRODUCTION 

The  Rhinotragini  comprise  a  large  group  of 
Neotropical  Cerambycidae  which  is  particularly 
abundant  and  diverse  in  South  America.  The 
group  is  of  special  interest  because  of  the  re- 
markable degree  of  mimicry  exhibited  by  most 
species.  This  is  evident  in  modifications  in  form, 
coloration,  and  behavior,  with  the  various  taxa 
utilizing  a  wide  range  of  models,  particularly 
bees,  wasps,  and  "protected"  beetles  of  other 
families.  Frequently,  the  mimicry  is  expressed 
in  strong  sexual  dimorphism  and  dichromatism 
with  males  and  females  utilizing  different 
models. 

The  number  of  species  diminish  as  the  group 
extends  northward  from  South  America,  and 
only  one  is  thus  far  known  to  occur  in  the  United 
States.  As  presently  defined,  the  Mexican  fauna 
consists  of  22  species  in  7  genera.  Some  of  the 
species  appear  to  be  restricted  to  Mexico,  while 
others  range  into  Central  America  to  Costa  Rica 
or  Panama.  One  is  also  found  in  South  America. 

Tribe  RHINOTRAGINI  Lacordaire 

Rhinotragides  LACORDAIRE,  1869:497. 
Rhinotraginae  BATES,  1873:21. 
Rhinotragini  LINSLEY,  1963:164. 


Body  more  or  less  elongate.  Head  prolonged 
below  eyes  to  form  a  muzzle;  antennae  usually 
shorter  than  body  in  both  sexes,  outer  segments 
more  or  less  serrated;  eyes  finely  faceted,  with 
lower  lobes  very  large,  nearly  contiguous  in 
front  in  males;  palpi  short,  last  segment  subcy- 
lindrical  or  ovate-cylindrical,  apex  truncate; 
mandibles  short.  Pronotum  cylindrical  or  round- 
ed, without  a  lateral  spine  or  tubercle;  proster- 
num  with  intercoxal  process  narrow,  level,  an- 
terior coxae  usually  obliquely  exserted,  cavities 
feebly  angulate  externally:  intermediate  coxal 
cavities  open  externally;  episterna  of  metatho- 
rax  very  large,  triangular,  and  in  front  broad  and 
narrowly  separated  from  coxae.  Elytra  nearly 
always  modified,  attenuated,  subulate,  or 
squamiform. 

KEY  TO  THE  MEXICAN  GENERA  OF 
RHINOTRAGINI 

1.        Elytra  short,  not  extended  over  abdo- 
men        2 

Elytra  longer,  extended  at  least  partially 
over  abdomen 4 

2(1).   Elytra  narrowing  posteriorly,  apices  de- 
hiscent and  narrowly  rounded  3 


[69] 


70 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  3 


Elytra  not  narrowing  posteriorly,  apices 

obliquely  truncate  Tomopterus 

3(2).  Pronotum  with  disk  punctate,  calluses, 
if  present,  small,  rounded;  posterior 
tibiae  with  very  dense  brush  of  long 
pubescence  Epimelitta 

Pronotum  with  three  longitudinal,  gla- 
brous calluses  on  disk;  posterior  tibi- 
ae with  apical  brush  small,  the  pubes- 
cence short  Bromiades 

4(1).  Pronotum  uniformly  punctate  or  with 
rounded  calluses  on  disk;  femora 
moderately  clavate 6 

Pronotum  usually  with  three  distinct, 
longitudinal  calluses  on  disk;  anterior 
and  intermediate  femora  often  very 

strongly  clavate 5 

5(4).  Antennae  short,  serrate  apically;  ante- 
rior and  intermediate  femora  very 
strongly  clavate Acyphoderes 

Antennae  slender,  elongate,  filiform  api- 
cally; anterior  and  intermediate  fem- 
ora narrowly  clavate  ____  Stenopseutes 
6(4).  Elytra  with  disk  vitreous  transparent; 
pronotum  about  as  long  as  broad  or 
slightly  longer Odontocera 

Elytra  with  disk  shining,  not  transpar- 
ent; pronotum  narrow,  distinctly 
longer  than  broad  Ommata 

Genus  Tomopterus  Audinet-Serville 

Tomopterus  AUDINET-SERVILLE,  1833:544;  NEWMAN  1840:21: 
WHITE  1855:176;  THOMSON  1860:166,  168;  1864:163,  416; 
LACORDAIRE  1869:509;  BATES  1870:329:  1880:44. 

This  genus  may  be  recognized  by  the  short, 
rather  squat  form,  short,  subserrate  antennae, 
rounded  pronotum,  short,  apically  truncate  ely- 
tra, and  arcuate  posterior  femora. 

TYPE-SPECIES:  Tomopterus  staphylinus  Au- 
dinet-Serville, 1833  (monotypic). 

The  species  of  Tomopterus  appear  to  mimic 
eumenine  vespids.  Two  species  occur  in  Mexi- 
co. 

Tomopterus  vespoides  White 

Tomopterus  vespoides  WHITE,  1855:176,  pi.  5,  fig.  8;  BATES 
1880:44;  1885:291;  FISHER  1930:17. 

MALE:  Integument  black,  antennae  reddish 
distally,  apices  of  hind  femora  and  tibiae  pale, 
elytra  usually  narrowly  reddish  along  lateral 
margins  and  with  a  reddish  median  stripe  ex- 
tending from  humeri  arcuately  to  middle  of 


apex.  Head  with  eyes  contiguous  on  front;  an- 
tennae with  segments  from  fifth  expanded,  sub- 
serrate.  Pronotum  almost  as  broad  as  elytra; 
disk  coarsely,  contiguously  punctate;  apex  and 
base  with  bands  of  yellow,  appressed  pubes- 
cence. Scutellum  elongate,  triangular,  nonpu- 
bescent.  Elytra  moderately  coarsely,  densely 
punctate,  sparsely  pubescent.  Abdomen  with 
sternites  margined  with  yellowish,  appressed 
pubescence;  parameres  prominent.  Length,  8- 
10  mm. 

FEMALE:  Form  more  robust.  Head  with  eyes 
separated  on  front  by  about  width  of  antennal 
scape.  Scutellum  yellow-pubescent.  Elytra 
black  medially,  remainder  reddish.  Abdomen 
bulbous,  pointed  apically.  Length,  8-10  mm. 

TYPE-LOCALITY:  Guatemala. 

RANGE:  Tamaulipas,  Mexico  to  Panama. 

FLOWER  RECORDS  (Costa  Rica):  Bixa,  For- 
steronia,  Byrsonima,  Paullinia,  Cor  dm. 

NEW  RECORDS:  1  female,  10  miles  [16  km]  N  Cuidad  Vic- 
toria, Tamaulipas,  20  Aug.  1941  (H.  S.  Dybas);  1  female,  9 
miles  [ca.  14  km]  S  Tuxpan,  Veracruz,  11  June  1961  (Mich- 
ener  and  Ordway);  1  female,  Tehuantepec,  Oaxaca,  8  July 
1962. 

Tomopterus  exilis,  new  species 

(Figure  1) 

FEMALE:  Form  small,  rather  slender;  integu- 
ment black,  antennae  brownish;  elytra  narrowly 
pale  at  base,  each  side  with  a  broad,  oblique, 
pale  vitta  extending  from  behind  humerus  to 
apex;  anterior  and  intermediate  legs  brownish, 
posterior  femora  pale  over  basal  one-half;  pu- 
bescence silvery,  appressed.  Head  with  front 
rather  short,  moderately  coarsely,  densely 
punctate;  eyes  separated  on  front  by  little  more 
than  diameter  of  antennal  scape;  separated  on 
vertex  by  about  twice  diameter  of  scape:  anten- 
nae clavate,  enlarging  from  fifth  segment,  outer 
segments  broader  than  long,  with  small  apical 
poriferous  areas,  third  segment  longer  than  first, 
fourth  shorter  than  first,  fifth  longer  than  fourth, 
basal  segments  with  a  few,  long,  erect  setae  be- 
neath. Pronotum  as  long  as  broad,  cylindrical; 
disk  coarsely,  deeply  reticulate-punctate;  apex 
narrowly  impressed,  base  rather  broadly  im- 
pressed; apex  and  base  with  narrow  transverse 
silvery  bands  of  appressed  pubescence,  each 
side  with  a  narrower  band  behind  middle  ex- 
tending from  sides  to  coxal  cavities;  long,  pale, 
erect  hairs  numerous;  prosternum  densely  punc- 
tate, sparsely  clothed  with  long,  erect  hairs; 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


71 


mesosternum  with  a  broad  pubescent  band  over 
epimeron;  metasternum  sparsely  punctate,  with 
a  long  erect  seta  rising  from  each  puncture,  a 
narrow,  oblique,  pubescent  band  present  pos- 
teriorly. Scutellum  densely  silver-pubescent. 
Elytra  broader  than  long,  densely,  deeply,  con- 
tiguously punctate;  pubescence  long,  erect;  pale 
vittae  in  form  of  a  broad  V;  apices  obliquely 
truncate.  Legs  slender;  femora  enlarged  over 
apical  one-half,  hind  pair  arcuate,  extending  al- 
most to  apex  of  abdomen;  tibiae  slender.  Ab- 
domen only  slightly  enlarged  posteriorly;  punc- 
tures and  pubescence  fine,  sternites  more 
densely  pubescent  along  posterior  margins;  last 
sternite  rounded  at  apex.  Length,  8  mm. 

Holotype,  female  (National  Museum  of  Natural  History) 
from  Canciin,  Quintana  Roo,  Mexico,  25  Apr.  1974  (D. 
Pletsch). 

The  more  slender  form,  cylindrical  rather 
than  transverse  pronotum,  and  longer  antennae 
will  separate  this  species  from  T.  vespoides 
White. 

Genus  Epimelitta  Bates 

Charts  NEWMAN,  1840:21;  THOMSON  1864:163;  LACORDAIRE 
1869:507  (name  preoccupied).  Type-species:  Charts  eu- 
phrosyne  Newman,  1840,  Thomson  designation,  1864). 

Epimelitta  BATES,  1870:330;  1873:123. 

Charisia  CHAMPION,  1892:161  (new  name  for  Charts  New- 
man, 1840,  preoccupied). 

The  Mexican  species  in  this  genus  may  be  rec- 
ognized by  the  short  elytra,  lack  of  longitudinal 
calluses  on  the  pronotum,  and  densely  tufted 
posterior  tibiae. 

TYPE-SPECIES:  Epimelitta  meliponica  Bates, 
1870  (by  present  designation). 

Most  species  of  Epimelitta  mimic  meliponid 
bees.  The  group  is  dominantly  South  American 
with  only  two  species  presently  known  from 
Mexico. 

Epimelitta  nigerrima  (Bates) 

Charisia  nigerrima  BATES,  1892:160,  pi. 6,  fig.  8. 
Epimelitta  nigerrima;  AURIVILLIUS  1912:284. 

FEMALE:  Integument  black,  antennae  reddish 
brown.  Antennae  short,  serrate  from  fifth  seg- 
ment. Pronotum  densely  clothed  with  long  erect 
pubescence;  disk  callused  behind  middle.  Elytra 
strongly  dehiscent,  densely  punctate.  Legs 
short,  femora  and  tibiae  with  dense  tufts  of  dark 
pubescence.  Length,  11  mm. 

TYPE-LOCALITY:  Atoyac,  Veracruz,  Mexico. 

Only  the  holotype  of  this  species  is  known. 


FIGURE  1.     Dorsal  view  of  pronotum  and  body  ofTomop- 
terus  exit  is  Chemsak  and  Linsley,  female. 


Epimelitta  aureopilis  Fisher 

Epimelitta  (?)  aureopilis  FISHER,  1953:16. 

MALE:  Form  slender,  elongate;  head,  pro- 
thorax,  part  of  metasternum  and  often  bases  of 
hind  femora  black,  antennae  reddish  brown, 
elytra  testaceous  except  for  narrow  black  lateral 
and  sutural  margins,  legs  and  abdomen  partially 
reddish,  posterior  tarsi  yellow.  Head  small, 
front  golden  pubescent;  antennae  slender,  mod- 
erately serrate.  Pronotum  densely,  coarsely 
punctate;  apex  and  base  with  a  broad  band  of 
appressed  golden  pubescence,  basal  band  reach- 
ing to  about  middle  at  center.  Elytra  dehiscent, 
sparsely  pubescent.  Legs  with  posterior  pair 
very  elongate,  femora  feebly  clavate,  tibiae  with 


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a  dense  tuft  of  long  reddish-orange  pubescence. 
Abdomen  elongate,  slender,  last  sternite  barely 
impressed.  Length,  16-21  mm. 

FEMALE:  Form  and  coloration  similar.  Ab- 
domen with  last  sternite  narrowly  rounded  at 
apex.  Length,  18-23  mm. 

TYPE-LOCALITY:  Tolome,  Veracruz,  Mexico. 

This  species  differs  greatly  in  facies  from  oth- 
er known  Epimelitta.  Its  long,  slender  form  with 
very  long  posterior  legs,  make  it  especially  dis- 
tinctive. However,  except  for  these  character- 
istics, aureopilis  is  structurally  similar  to  other 
species  within  the  genus. 

Fisher  (1953)  tentatively  assigned  aureopilis 
to  Epimelitta  because  of  the  difficulty  in  delim- 
iting genera  of  Rhinotragini.  We  have  retained 
it  in  this  genus  since  it  is  beyond  the  intent  of 
this  paper  to  attempt  to  clarify  the  generic  clas- 
sification of  this  primarily  South  American 
group. 

NEW  RECORDS:  3  females,  3  males,  Cotaxtla  Exp.  Sta., 
Cotaxtla,  Veracruz,  1  Aug.  1962  (D.  H.  Janzen);  1  male, 
Ixmal,  Yucatan  (G.  F.  Gaumer);  1  female,  30  miles  [48  km] 
NE  Tehuantepec,  Oaxaca,  8  July  1955  (D.  Giuliani). 

Genus  Bromiades  Thomson 

Bromiades   THOMSON,    1864:165;   LACORDAIRE    1869:506; 
BATES  1873:120;  ZAYAS  1975:130. 

The  tricallused  pronotum,  strongly  serrate  an- 
tennae, and  short,  tapered  elytra  distinguish  this 
genus. 

TYPE-SPECIES:  Odontocera  brachyptera 
Chevrolat,  1838  (by  original  designation). 

Bromiades  brachyptera  (Chevrolat) 

Odontocera  brachyptera  CHEVROLAT,   1838:285;  JACQUELIN 

ouVAL  in  SACRA  1857:269,  pi.  10,  fig.  9. 
Bromiades  brachyptera:  THOMSON  1864:165;  BATES  1873:120; 

FISHER  1930:13;  LINSLEY   1935:85,  pi.  2,  fig.  3;  ZAYAS 

1956:113;  1975:131,  pi.  17,  fig.  a. 
Bromiades  meridionalis  FISHER,   1930: 14  (type-locality,  Ca- 

bima,  Panama).  NEW  SYNONYMY. 

MALE:  Integument  black,  shining,  antennal 
segments  four  to  usually  eight,  yellowish  basal- 
ly;  elytra  broadly  testaceous  medially;  femora 
usually  with  basal  one-half  testaceous;  anterior 
and  intermediate  tibiae  basally  and  posterior 
pair  except  apically,  testaceous.  Pronotum 
densely  clothed  with  golden,  recumbent  pubes- 
cence between  calluses.  Abdomen  not  modified 
apically.  Length,  15-17  mm. 

FEMALE:  Form  and  coloration  similar  to  male, 


abdomen  broader.  Antennae  occasionally  al- 
most all  black.  Length,  16-18  mm. 

TYPE-LOCALITY:  Cuba. 

RANGE:  Central  Mexico  to  Colombia  and 
Cuba. 

HOST  PLANTS:  Andira  inermis  (W.  Wright) 
H.B.K.  ex  D.C.  (Cuba). 

FLOWER  RECORDS.  Costa  Rica:  Cordia,  Ca- 
searia,  Forsteronia,  Coccoloba,  asclepiad  vine. 
Mexico:  Spondias. 

Color  variation  is  apparent  in  the  antennae 
and  legs.  Specimens  from  Panama  have  domi- 
nantly  black  antennae  and  orange  posterior  fem- 
ora and  tibial  brushes.  This  type  of  color  vari- 
ation occurs  in  numerous  species  throughout  the 
tribe. 

We  have  seen  no  additional  specimens  from 
Mexico  since  those  reported  by  Linsley  (1935) 
from  Bejucos. 

Genus  Acyphoderes  Audinet-Serville 

Acyphoderes  AUDINET-SERVILLE,  1833:549;  WHITE  1855:194; 
THOMSON  1860:179;  1864:165;  LACORDAIRE  1869:505; 
BATES  1873:117;  1880:43. 

This  genus  may  be  recognized  by  the  dorsal 
callosities  of  the  pronotum.  The  antennae  are 
distally  serrate  and  usually  extend  to  about  the 
middle  of  the  elytra.  The  elytra  are  subulate  or 
elongate,  narrowed  posteriorly  and  extend  to  the 
abdomen.  The  anterior  and  intermediate  femora 
are  usually  strongly  clavate.  Males  of  most 
species  have  the  last  abdominal  sternite  modi- 
fied in  varying  degrees. 

TYPE-SPECIES:  Acyphoderes  aurulenta  Kirby, 
1818  (Thomson  designation,  1864). 

Acyphoderes,  with  ten  species,  is  the  largest 
group  of  Rhinotragini  found  in  Mexico.  Two 
other  species,  velutinus  Bates,  1885,  and  ves- 
piventris  Bates,  1880,  described  originally  from 
Guatemala,  probably  also  occur  in  Mexico. 

Key  to  the  Mexican  species  of  Acyphoderes 

\.  Pronotum  black,  with  discal  callosities 
shining,  punctures  sparse  or  absent  __ 

2 

Pronotum  reddish  or  infuscated,  discal 
callosities  opaque,  densely  punctate 
5 

2(1).  Pronotum  with  transverse  or  longitudi- 
nal bands  of  dense,  appressed  pubes- 
cence   3 

Pronotum  rather  sparsely  pubescent, 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


73 


without  basal  and  apical  transverse 
pubescent  bands  4 

3(2).  Pronotum  densely  pubescent  along  api- 
cal and  basal  margins,  dorsal  calluses 
usually  punctate,  lateral  pair  rounded, 
punctures  very  dense,  confluent. 
Length,  13-20  mm.  Sinaloa  to  Oaxaca 

and  Veracruz cribricollis 

Pronotum  densely  pubescent  except  on 
calluses,  calluses  shining,  elongate, 
not  punctate,  punctures  obscured. 
Length,  14-18  mm.  Sinaloa  to  Chia- 
pas    amoena 

4(2).  Elytra  subulate,  extending  only  to  an- 
terior margin  of  second  abdominal 
segment;  abdomen  strongly  inflated 
toward  apex.  Length,  17-21  mm. 

Nayarit  to  Oaxaca  and  Veracruz 

sexualis 

Elytra  more  gradually  tapering,  extend- 
ing over  first  three  abdominal  seg- 
ments; abdomen  narrowed  toward 
apex.  Length,  14-18  mm.  Veracruz  to 
Costa  Rica  fulgida 

5(1).  Elytra  with  apices  rounded,  as  broad  as 
or  broader  than  diameter  of  antennal 

scape 6 

Elytra  with  apices  truncate  to  emargin- 
ate,  narrower  than  diameter  of  anten- 
nal scape  7 

6(5).  Pronotum  deeply  impressed  at  base  and 
apex,  with  bands  of  dense  appressed 
pubescence  in  impressions;  elytra 
black  around  scutellum.  Length,  20- 

25  mm.  Colima  prolixa 

Pronotum  not  deeply  and  broadly  im- 
pressed at  base  and  apex,  without 
transverse  bands  of  dense  pubes- 
cence; elytra  pale  around  scutellum. 
Length,  11-18  mm.  Baja  California  __ 
delicata 

7(5).  Pronotum  with  apical  and  basal  trans- 
verse bands  of  dense,  appressed  pu- 
bescence, dorsal  calluses  moderately 

elevated 8 

Pronotum  rather  uniformly  pubescent, 
without  transverse  bands,  dorsal  cal- 
luses prominently  elevated.  Length, 
15-20  mm.  Colima  to  Oaxaca  and  Ve- 
racruz    acutipennis 

8(7).  Elytra  2.5  or  more  times  longer  than 
basal  width,  extending  at  least  to  third 
abdominal  segment  9 


Elytra  short,  2.0  times  as  long  as  basal 
width,  extending  only  to  second  ab- 
dominal segment.  Length,  10-18  mm. 
Campeche  to  Honduras  ..  _  yucateca 
9(8).  Legs  with  hind  femora  pedunculate;  ab- 
domen abruptly  inflated  toward  apex; 
elytra  with  two  oblique,  usually 
brownish,  scabrous  bands  at  base. 
Length,  11-16  mm.  Sinaloa  to  Nay- 
arit    parva 

Legs  with  hind  femora  gradually  en- 
larged; abdomen  gradually  expanded 
toward  apex;  elytra  punctate  at  base, 
with  a  triangular  yellowish  area 
around  scutellum.  Length,  13-22  mm. 
Sonora  and  Tamaulipas  to  Costa  Rica 

__  suavis 


Acyphoderes  cribricollis  Bates 

Acyphoderes  cribricollis  BATES,  1892:160,  pi.  6,  fig.  6;  LINS- 
LEY 1935:84;  CHEMSAK  1967:74  (lectotype). 

MALE:  Form  slender,  elongate;  integument 
black;  antennae  reddish  brown,  usually  infus- 
cated  apically;  elytra  testaceous,  shining,  mar- 
gins dark,  legs  pale,  hind  pair  with  femoral  club 
and  apices  of  tibiae  brownish;  abdomen  with 
first  two  segments  yellowish,  usually  infuscated 
apically,  other  segments  usually  reddish.  Head 
with  front  densely  pubescent,  eyes  separated  by 
less  than  diameter  of  antennal  scape.  Pronotum 
longer  than  broad;  apex  and  base  broadly  im- 
pressed; disk  with  median  callus  punctate,  lat- 
eral pair  more  elevated  anteriorly,  glabrous, 
rounded;  punctures  dense,  opaque;  apical  and 
basal  margins  with  broad  bands  of  dense,  ap- 
pressed, golden  pubescence,  bands  joined  lat- 
erally and  usually  extended  longitudinally  on 
disk  on  each  side  of  middle;  pro-  and  mesoster- 
num  pubescent;  anterior  one-half  of  metaster- 
num  and  posterior  end  of  episternum  densely 
pubescent.  Elytra  extending  to  posterior  margin 
of  third  abdominal  segment;  base  and  disk  fine- 
ly, sparsely  punctate,  margins  densely  punctate; 
apices  rounded,  about  as  broad  as  antennal 
scape.  Legs  with  femora  clavate,  anterior  and 
intermediate  pairs  moderately  clavate.  Abdo- 
men slender,  slightly  enlarged  toward  apex;  last 
sternite  deeply,  broadly  excavated,  margins 
prominent,  elevated  apically.  Length,  14-20 
mm. 

FEMALE:  Form  similar.  Head  with  eyes 
widely  separated  on  front.  Abdomen  with  last 
sternite  not  modified.  Length,  13-20  mm. 


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FIGURE  2.     Known  distribution  of  Acyphoderes  cribricollis  Bates. 


TYPE-LOCALITY:  Ventanas,  Durango. 

FLOWER  RECORDS:  Croton,  Xanthoxylum, 
Jatropha. 

The  dense  pubescence,  rounded,  glabrous  cal- 
luses, and  dense,  opaque  punctation  of  the 
pronotum  distinguish  this  species. 

NEW  RECORDS:  See  Figure  2. 

Acyphoderes  amoena,  new  species 

(Figure  3) 

MALE:  Form  moderate  sized;  integument 
black;  antennae  reddish  brown;  elytra  shining, 
yellow-brown,  margins  black,  base  often  with 
two  short  dark  vittae  extending  back  from  inside 
of  humeri;  legs  orange-brown,  posterior  femora 
often  dark  over  clavate  portion;  abdomen  usu- 
ally pale  and  dark.  Head  with  eyes  separated  on 
front  by  less  than  diameter  of  antennal  scape; 
front  and  vertex  densely  clothed  with  appressed 
golden  pubescence.  Pronotum  longer  than 
broad,  sides  impressed  before  and  behind  mid- 
dle; disk  uneven,  dorsal  calluses  glabrous,  ele- 
vated, sides  with  smaller  calluses  before  middle, 
pubescence  between  calluses  dense,  appressed, 
golden,  obscuring  surface,  long,  erect  hairs  rath- 


er sparse;  prosternum  densely  pubescent;  meso- 
sternum  densely  clothed  with  appressed  golden 
pubescence;  metasternum  densely  pubescent 
except  for  rectangular  areas  near  base,  epister- 
num  with  dense  pubescent  patches  posteriorly 
and  anteriorly  on  upper  margin.  Elytra  elongate, 
extending  to  fourth  abdominal  segment;  lateral 
margins  darkened,  narrowly  punctate,  sutural 
margins  elevated,  narrowly  dark;  disk  shining, 
very  sparsely  punctate  except  on  base;  apices 
rounded,  broad.  Legs  with  anterior  and  inter- 
mediate femora  moderately  clavate;  posterior 
femora  gradually  clavate;  posterior  tibiae  dense- 
ly pubescent  over  apical  one-half.  Abdomen  not 
apically  enlarged;  sternites  with  small  patches 
of  dense  pubescence  at  sides  of  apical  margins; 
last  sternite  broadly  excavated,  margins  elevat- 
ed toward  apex,  angles  blunt.  Length,  14-18 
mm. 

FEMALE:  Form  similar.  Head  with  eyes 
widely  separated  on  front.  Abdomen  with  last 
sternite  broadly  rounded  at  apex.  Length  16-18 
mm. 

Holotype,  male  (California  Academy  of  Sciences)  from  3 
miles  [ca.  5  km]  E  Villa  Union,  Sinaloa,  Mexico,  24  July  1972, 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


75 


FIGURE  3.    Acyphoderes  amoena  Chemsak  and  Linsley,  male. 


on  Jatropha  curcas  flowers  (J.  A.  and  M.  A.  Chemsak,  A.  E. 
and  M.  M.  Michelbacher).  Paratypes  as  follow:  1  male,  same 
data  as  type;  1  female,  16  miles  [ca.  26  km]  NE  San  Bias, 
Nayarit,  21  July  1963  (R.  Westcott);  1  male,  14  miles  [ca.  23 


km)  NW  Tehuantepec,  Oaxaca,  26  June  1961,  on  flowers  of 
Croton  (Univ.  Kans.  Mex.  Exped.);  1  male,  23  miles  [ca.  37 
km]  S  Mati'as  Romero,  Oaxaca,  14  Aug.  1963  (Parker  and 
Stange);  1  female,  Playa  La  Ventosa,  3.5  miles  [ca.  5.6  km] 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  3 


NE  Salina  Cruz,  Oaxaca,  28  July  1970  (Fisher,  Sullivan);  1 
female,  5  miles  [ca.  8  km]  NW  Escuinapa,  Sinaloa,  25  July 
1971  (Fisher). 

We  are  also  assigning  to  this  species  five  spec- 
imens from  various  localities  in  Mexico  which 
differ  in  having  the  antennae  black,  with  seg- 
ments 8-11  all  or  partially  yellow.  In  most  of 
these,  the  two  basal  black  vittae  of  the  elytra 
are  very  distinct  and  the  pubescence  is  more 
silvery  than  golden.  Structurally  they  are  iden- 
tical with  the  typical  forms. 

Records  are:  1  female,  Tecolopa,  Colima,  31  July  1954  (Ca- 
zier,  Gertsch,  Bradts);  1  female,  10  miles  [ca.  16  km]  W  Col- 
ima, 1  Aug.  1954  (Cazier,  Gertsch,  Bradts);  1  male,  Manzan- 
illo,  Colima,  12  July  1956  (R.  and  K.  Dreisbach);  1  male, 
Acapulco,  Guerrero,  30  July  1933  (M.  A.  Embury);  1  male,  31 
miles  [ca.  50  km]  SE  Comitan,  Chiapas,  18  June  1965  (Burke, 
Meyer,  Schaffner). 

Acyphoderes  sexualis  Linsley 

Acyphoderes  sexualis  LlNSLEY,  1934:349;  1935:84,  pi.  2,  figs. 
1,  la. 

MALE:  Form  rather  elongate;  integument 
black,  antennae  with  basal  segments  infuscated 
beneath,  reddish  above,  outer  segments  pale  an- 
nulate basally;  elytra  with  disk  pale  vitreous; 
abdomen  with  first  segment  reddish  at  middle; 
posterior  legs  with  femora  narrowly  pale  basal- 
ly, tibiae  yellowish  at  basal  one-half.  Head  with 
eyes  separated  on  front  by  about  diameter  of 
third  antennal  segment.  Pronotum  slightly 
broader  than  long,  sides  impressed  behind  mid- 
dle, base  narrower  than  apex;  disk  with  three 
longitudinal,  sparsely  punctate,  shining  calluses; 
punctures  between  calluses  fine,  dense;  pubes- 
cence fine,  moderately  dense,  appressed,  long 
erect  hairs  numerous;  mesosternum  with  epi- 
meron  densely  pubescent;  metasternum  with  a 
broad,  densely  pubescent  band  anteriorly  and  a 
narrow  band  along  posterior  margin.  Elytra  sub- 
ulate, extending  to  anterior  portion  of  second 
abdominal  segment;  lateral  margins  broadly 
black  and  densely  punctate,  sutural  margins  nar- 
rowly black,  impunctate;  base  scabrous,  with 
two  oblique  black  vittae  behind  scutellum;  disk 
vitreous,  almost  impunctate.  Legs  with  anterior 
and  intermediate  femora  strongly  clavate;  pos- 
terior tibiae  with  a  dense  brush  of  hairs  over 
apical  one-half.  Abdomen  strongly  inflated  to- 
ward apex;  sternites  with  first  three  segments 
margined  at  sides  and  apices  with  dense  pubes- 
cence; last  sternite  with  a  large  circular  plate, 


sides  prominently  elevated;  claspers  large. 
Length,  20-21  mm. 

FEMALE:  Form  similar.  Head  with  eyes 
widely  separated  on  front.  Pronotum  with  dorsal 
calluses  less  prominent.  Abdomen  with  last  ster- 
nite excavated  but  lacking  a  plate.  Length,  17 
mm. 

TYPE-LOCALITY:  Bejucos,  Temascaltepec, 
Mexico. 

FLOWER  RECORDS:  Acacia,  Spondias. 

NEW  RECORDS:  1  male,  Arroyo  Santiago,  near  Jesus  Maria, 
Nayarit,  5  July  1955  (B.  Malkin);  1  male,  Mexcala,  Guerrero, 
29  June  1951  (H.  Evans);  2  males,  20  miles  [32  km]  S  Matias 
Romero,  Oaxaca,  25  June  1961,  on  flowers  of  Acacia  (Univ. 
Kans.  Mex.  Exped.);  1  female,  8  miles  [ca.  13  km]  NE  Ca- 
temaco,  Veracruz,  3  July  1971  (Clark,  Murray,  Hart,  Schaff- 
ner). 

Acyphoderes  fulgida,  new  species 

(Figure  4) 

MALE:  Form  moderate  sized,  tapering;  integ- 
ument black,  shining;  antennae  often  brownish 
toward  apex;  elytra  with  disk  testaceous,  vitre- 
ous; hind  femora  usually  testaceous  over  basal 
one-half  except  at  extreme  base.  Head  with  eyes 
contiguous  on  front;  inner  margins  of  front  ele- 
vated, nonpubescent.  Pronotum  longer  than 
broad,  sides  impressed  behind  middle;  disk  with 
dorsal  calluses  elongate,  barely  punctate,  shin- 
ing; sides  with  oblique  calluses  on  anterior  one- 
half;  punctures  between  calluses  fine,  dense; 
pubescence  fine,  pale,  appressed,  denser  on 
sides,  long,  pale,  erect  hairs  numerous;  proster- 
num  densely  punctate  at  middle,  densely  clothed 
with  long,  pale,  erect  hairs;  mesosternum  with 
a  dense,  pale,  pubescent  patch  on  epimeron; 
metasternum  densely  pubescent  over  apical  one- 
fourth,  episternum  with  a  small  pubescent  patch 
at  end.  Elytra  elongate,  extending  to  fourth  ab- 
dominal segment;  lateral  margins  broadly  black 
at  base,  narrowly  black  toward  apex,  sutural 
margins  more  broadly  black  at  base;  punctures 
dense  on  dark  margins,  disk  very  sparsely  punc- 
tate; apices  rounded,  broader  than  antennal 
scape.  Legs  with  anterior  and  intermediate  fem- 
ora strongly  clavate,  clavae  large;  posterior  fem- 
ora gradually  clavate;  posterior  tibiae  with  a 
moderately  dense  brush  of  dark  hairs  over  apical 
one-half.  Abdomen  not  enlarged  apically;  ster- 
nites one  to  three  densely  clothed  with  pale  pu- 
bescence along  lateral  and  apical  margins  and 
with  a  double  row  at  middle;  last  sternite  shal- 
lowly  excavated  with  a  large  blunt  tubercle  on 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


77 


FIGURES  4-5.     Dorsal  view  of  pronotum  and  body  of  (left)  Acyphoderes  fulgida  Chemsak  and  Linsley,  male,  and  (right) 
Acyphoderes  parva  Chemsak  and  Linsley,  male. 


each  side  at  base,  claspers  densely  pubescent. 
Length,  16-18  mm. 

FEMALE:  Form  similar.  Head  with  eyes  sep- 
arated on  front  by  more  than  diameter  of  anten- 
nal  scape.  Pronotum  very  sparsely  punctate  be- 
tween calluses.  Abdomen  with  last  sternite 
narrowly  rounded  at  apex.  Length,  14-18  mm. 

Holotype,  male;  allot)  pt  (California  Academy  of  Sciences) 
from  La  Pacifica,  4  km  NW  Canas,  Guanacaste  Prov.,  Costa 
Rica,  25  May  1974,  on  Casearia  nitida  flowers  (P.  Opler).  Six 
paratypes  (3  males,  3  females)  same  locality,  all  collected  by 
P.  Opler,  25  May  1974,  2-4  June  1973,  on  asclepid  vine,  3 
June  1971,  on  Trigonia  floribunda :  1  female,  2  miles  [ca.  3.2 
km]  NW  Canas,  18-21  May  1974  (E.  Giesbert);  2  females, 
Playa  Hermosa,  Guanacaste,  6-10  June  1974  (Giesbert). 


Two  additional  specimens  (1  male,  1  female)  from  Salto 
Eyiplantla,  near  Catemaco,  Veracruz,  Mexico,  17  Aug.  1976 
(J.  Hafernik  and  R.  Garrison)  are  also  assigned  to  this  species. 

In  addition  to  the  marked  sexual  dimorphism, 
this  species  varies  slightly  in  coloration.  Males 
occasionally  have  the  posterior  femora  yellow 
and  brownish.  Some  of  the  females  have  the 
femora  and  tibiae  pale  brownish  except  for  the 
apical  one-half  of  the  hind  tibiae. 

The  elongate  elytra,  punctation  of  the  prono- 
tum, and  nonbulbous  abdomen  will  separate  ful- 
gida from  sexualis.  The  structure  of  the  male 
copulatory  apparatus  is  also  strikingly  different 
in  the  two  species. 


78 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  3 


Acyphoderes  prolixa,  new  species 

MALE:  Form  very  elongate;  head  reddish;  an- 
tennae brownish;  pronotum  black;  dorsal  callus- 
es reddish;  elytra  testaceous,  margins  brownish, 
base  with  a  triangular  black  spot  around  scutel- 
lum;  thoracic  sterna  black;  legs  pale,  tibiae  dark 
on  dorsal  edge,  femora  black  internally  on  an- 
terior portion  of  club;  abdomen  pale  basally, 
reddish  toward  apex,  sternites  variably  infus- 
cated  at  bases.  Head  with  eyes  separated  on 
front  by  about  diameter  of  antennal  scape;  inner 
margins  of  front  densely  pubescent;  antennae 
with  segments  from  fourth  opaque.  Pronotum 
slightly  longer  than  broad,  sides  rounded;  apex 
and  base  broadly  impressed;  disk  convex,  dorsal 
calluses  moderately  elevated;  calluses  punctate, 
other  punctures  fine,  dense;  apex  and  base  with 
densely  pubescent  bands;  prosternum  densely 
pubescent;  mesosternum  with  epimeron  densely 
clothed  with  pale  appressed  pubescence;  meta- 
sternum  densely  pubescent  anteriorly,  epister- 
num  with  a  dense  pubescent  patch  posteriorly. 
Elytra  narrow,  extending  at  least  to  middle  of 
third  abdominal  segment;  lateral  margins  brown- 
ish, densely  punctate,  sutural  margins  elevated, 
pale  brownish;  disk  shining,  testaceous,  very 
sparsely  punctate;  apices  rounded,  about  as 
wide  as  antennal  scape.  Legs  with  anterior  and 
intermediate  femora  strongly  clavate;  posterior 
femora  petiolate,  clavae  moderate.  Abdomen 
elongate,  not  enlarged  apically;  sternites  densely 
punctate,  moderately  densely  pubescent;  fourth 
sternite  shallowly  impressed  at  apex;  last  ster- 
nite  broadly,  deeply  impressed,  margins  elevat- 
ed basally,  not  forming  angle  at  apex.  Length, 
21-25  mm. 

FEMALE:  Form  more  robust,  less  elongate. 
Head  with  eyes  widely  separated  on  front. 
Pronotum  reddish,  black  on  apical  and  basal 
margins;  metepisternum  partially  reddish.  Ab- 
domen enlarged  apically;  last  sternite  not  im- 
pressed. Length,  20-24  mm. 

Holotype,  male,  allotype  (American  Museum  of  Natural  His- 
tory) and  6  paratypes  (3  males,  3  females)  from  10  miles  [ca. 
16  km]  W  Colima,  Colima,  Mexico,  1  Aug.  1954  (M.  Cazier, 
W.  Gertsch,  Bradts). 

This  species  may  be  recognized  by  the  elon- 
gate form  and  black  triangular  patch  around  the 
scutellum.  The  coloration,  more  petiolate  pos- 
terior femora,  and  broadly  excavated,  margined 
last  abdominal  sternite  of  the  males  will  separate 
prolixa  from  suavis. 


Acyphoderes  delicata  Horn 

Acyphoderes  delkatus  HORN,  1894:400;  HAMILTON  18%:  168; 
LINSLEY  1942:54. 

MALE:  Form  slender,  gradually  tapering;  in- 
tegument reddish;  antennae  orange-brown;  ely- 
tra shining  pale  brownish,  margins  darker;  un- 
derside partially  infuscated;  legs  pale,  femora 
with  clavate  portions  brownish,  hind  tibiae 
broadly  brown  at  apices.  Head  with  eyes  sepa- 
rated on  front  by  more  than  diameter  of  antennal 
scape.  Pronotum  as  long  as  broad,  broadly 
rounded;  disk  opaque,  median  callus  narrow, 
elongate,  lateral  pair  broader,  arcuate,  densely 
punctate;  pubescence  fine,  uniform,  dense, 
short,  erect;  mesosternum  with  sides  densely 
pubescent;  metasternum  densely  pubescent 
over  anterior  one-half,  episternum  with  a  dense 
pubescent  patch  posteriorly.  Elytra  extending  to 
fourth  abdominal  segment;  base  narrowly  yel- 
lowish, sparsely  punctate,  margins  densely 
punctate,  disk  sparsely  punctate;  apices  round- 
ed, much  broader  than  antennal  scape.  Legs 
with  anterior  and  intermediate  femora  clavate, 
clavae  small;  posterior  tibiae  densely  pubescent 
at  apical  one-half.  Abdomen  slender,  not  en- 
larged apically;  last  sternite  very  shallowly  im- 
pressed at  apex,  sides  not  margined.  Length,  1 1- 
18  mm. 

FEMALE:  Not  seen. 

TYPE-LOCALITY:  El  Taste,  Baja  California 
Sur. 

This  species  is  distinctively  characterized  by 
the  inflated,  opaque  pronotum  and  apically  slen- 
der abdomen. 

NEW  RECORDS:  2  males,  Santa  Victoria  Trail:  La  Burrera- 
La  Laguna  ridge,  1,200-1,350  m,  Baja  California  Sur,  28  Aug. 
and  2  Sep.  1977  (R.  L.  Westcott). 

Acyphoderes  acutipennis  Thomson 

Acyphoderes  acutipennis  THOMSON,  1860:179;  BATES,  1880:43; 
1885:290. 

MALE:  Form  elongate,  strongly  tapered;  in- 
tegument brownish;  calluses  of  pronotum  often 
yellowish;  elytra  yellowish  brown.  Head  with 
eyes  separated  on  front  by  about  diameter  of 
antennal  scape;  pubescence  sparse.  Pronotum 
longer  than  broad,  sides  impressed  behind  mid- 
dle; disk  with  calluses  prominently  elevated, 
punctate,  subopaque;  pubescence  moderately 
dense,  pale,  appressed,  rather  uniform;  meso- 
sternum with  a  dense  patch  of  appressed  pubes- 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


79 


cence  at  sides;  metasternum  with  a  small  pu- 
bescent patch  on  posterior  part  of  episternum. 
Elytra  elongate,  strongly  tapered,  extending  to 
anterior  margin  of  third  abdominal  segment;  lat- 
eral margins  broadly  brownish,  densely  punc- 
tate, sutural  margins  narrowly  brownish;  base 
finely,  densely  asperate  punctate;  disk  finely, 
sparsely  punctate,  each  puncture  bearing  a  short 
erect  seta;  apices  truncate,  narrower  than  di- 
ameter of  antennal  scape.  Legs  with  anterior  and 
intermediate  femora  strongly  clavate.  Abdomen 
gradually  inflated  toward  apex;  last  sternite  shal- 
lowly  impressed  at  apex,  sides  barely  margined. 
Length,  15-20  mm. 

FEMALE:  Form  similar.  Head  with  eyes  more 
widely  separated  on  front.  Abdomen  more  bul- 
bous, last  sternite  not  impressed.  Length,  16-20 
mm. 

TYPE-LOCALITY:  Mexico. 

The  prominent,  often  pale  calluses  of  the 
pronotum  and  rather  elongate,  apically  narrow 
elytra  make  this  species  distinctive. 

New  records:  4  males,  2  females,  Temascal,  Oaxaca.  3  June 
1964,  1  Nov.  1963  (D.  Janzen):  1  male,  Tezonapa,  Veracruz, 
July  1906;  1  male,  Colima,  Colima  (Conradt). 

Acyphoderes  yucateca  (Bates),  new  combination 

Odontotera  yucate ca  BATES,  1892:159,  pi.  6,  fig.  7;  CHEMSAK 
1967:78  (lectotype). 

MALE:  Form  moderately  elongate,  strongly 
tapered;  integument  light  to  dark  reddish  brown; 
elytra  with  disk  testaceous.  Head  with  eyes  sep- 
arated on  front  by  about  diameter  of  antennal 
scape.  Pronotum  longer  than  broad,  sides  lightly 
impressed  behind  middle,  base  slightly  narrower 
than  apex;  disk  with  elongate  calluses  punctate, 
opaque;  punctation  dense,  shallow,  scabrous; 
base  and  apex  with  narrow  bands  of  dense,  ap- 
pressed  pubescence,  long  erect  hairs  rather 
sparse;  meso-  and  metasternum  with  small, 
densely  pubescent  patches  at  sides  of  posterior 
margins.  Elytra  strongly  tapered,  extending  to 
anterior  margin  of  second  abdominal  segment; 
lateral  margins  dark  from  behind  humeri  to  api- 
ces, densely  punctate,  sutural  margins  narrowly 
dark;  impunctate;  base  moderately  densely 
punctate,  punctures  sparse  toward  apex,  each 
bearing  a  short  seta;  humeri  and  often  a  V- 
shaped  basal  vitta  dark;  apices  truncate,  nar- 
rower than  diameter  of  antennal  scape.  Legs 
with  anterior  and  intermediate  femora  strongly 
clavate.  Abdomen  moderately  inflated  toward 


apex;  two  basal  segments  yellowish;  last  sternite 
shallowly  impressed,  sides  lightly  margined;  last 
tergite  densely  clothed  with  appressed  pubes- 
cence. Length,  10-18  mm. 

FEMALE:  Form  similar.  Head  with  eyes  more 
widely  separated  on  front.  Abdomen  more  in- 
flated toward  apex;  last  sternite  unmodified. 
Length,  12-18  mm. 

TYPE-LOCALITY:  Temax,  Yucatan,  Mexico. 

The  pubescent  bands  and  less  elevated  callus- 
es of  the  pronotum  will  distinguish  this  species 
from  acutipennis. 

NEW  RECORDS:  4  males.  Yucatan  (G.  F.  Gaumer);  3  males, 
3  females,  Piste,  Yucatan,  6  and  26  Sep.  1967  (E.  Welling);  1 
male,  Yaxche,  Yucatan,  10  Sep.  1964  (J.  and  D.  Pallister);  1 
male,  1  female,  X-Can,  Quintana  Roo,  25  June  1967,  26  Sep. 
1967  (Welling);  1  male,  Hwy.  180,  Quintana  Roo,  23  Mar.  1964 
(E.  L.  Mockford);  1  male,  26  miles  [ca.  42  km]  S  Yucatan 
line,  Campeche,  30  June  1963  (Porter);  1  male,  Campeche, 
Campeche,  30  June  1966  (McFadden);  1  male,  El  Zapotal,  2 
miles  [ca.  3.2  km]  S  Tuxtla  Gutierrez,  Chiapas,  9  July  1957 
(J.  Chemsak);  1  female,  Puerto  Castilla,  Honduras,  Sep.  1945 
(Crowell). 

Acyphoderes  parva,  new  species 

(Figure  5) 

MALE:  Form  rather  small,  short;  integument 
pale  brownish:  head,  pronotum,  underside,  and 
abdomen  variously  infuscated;  elytra  testa- 
ceous, dark  margined,  and  with  two  dark, 
oblique  basal  bands.  Head  with  eyes  separated 
on  front  by  a  little  more  than  diameter  of  anten- 
nal scape;  inner  margins  of  front  densely  pubes- 
cent. Pronotum  longer  than  broad,  sides  slightly 
impressed  behind  middle;  disk  convex,  dorsal 
calluses  rather  prominent,  punctate;  surface 
densely  punctate,  opaque;  apical  and  basal  mar- 
gins with  narrow  pubescent  bands;  sternum  with 
a  narrow,  densely  pubescent  band  at  sides; 
metasternum  densely  pubescent,  with  a  pubes- 
cent patch  at  end  of  episternum.  Elytra  extend- 
ing to  anterior  margin  of  third  abdominal  seg- 
ment; base  finely  scabrous,  densely  punctate; 
lateral  margins  densely  punctate,  disk  finely, 
sparsely  punctate;  apices  truncate,  narrower 
than  antennal  scape.  Legs  with  anterior  and  in- 
termediate femora  strongly  clavate,  clavae 
large;  hind  femora  petiolate,  clavae  short;  hind 
tibiae  arcuate.  Abdomen  enlarged  toward  apex; 
two  basal  segments  with  lateral  patches  of  dense 
pubescence;  last  sternite  broadly  excavated, 
margins  elevated  at  apex  forming  obtuse  angles. 
Length  11-15  mm. 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  3 


FEMALE:  Form  similar.  Head  with  eyes  more 
widely  separated  on  front.  Pronotum  with  dorsal 
calluses  less  prominent.  Abdomen  bulbous,  last 
sternite  not  impressed.  Length,  12-16  mm. 

Holotype,  male;  allotype  (California  Academy  of  Sciences) 
from  5  miles  [ca.  8  km]  N  Mazatlan,  Sinaloa,  Mexico,  1  Aug. 
1972,  on  flowers  of  Buddleia  wrightii  (J.  A.  and  M.  A.  Chem- 
sak),  9-15  Aug.  1970,  on  B.  wrightii  (J.  A.  Chemsak).  Para- 
types  as  follow:  1  female,  5  miles  [ca.  8  km]  N  Mazatlan,  9- 
15  Aug.  1970,  on  Buddleia  (J.  A.  Chemsak);  10  males,  Ma- 
zatlan, 28  Nov.  1962  (T.  Gantenbein);  1  female,  Acaponeta, 
Nayarit,  19  Nov.  1955  (E.  C.  Bay);  2  males,  15  miles  [ca.24 
km]  NW  Acaponeta,  19  Oct.  1964  (A.  E.  Michelbacher);  1 
male,  15  miles  [ca.  24  km]  S  Acaponeta,  20  Aug.  1964  (Burke 
and  Apperson);  1  female,  Teacapan,  Sinaloa,  29  June  1956  (R. 
and  K.  Dreisbach);  2  females,  Tuxpan,  Nayarit,  20  Aug.  1964 
(Burke  and  Apperson);  1  female,  60  miles  [ca.  97  km]  N  Tepic, 
Nayarit,  15  Aug.  1957  (J.  A.  Chemsak). 

This  species  averages  smaller  in  size  than 
most  other  Mexican  Acyphoderes.  It  can  be  sep- 
arated from  suavis  by  the  different  coloration, 
more  inflated  abdomen,  and  petiolate  posterior 
femora.  Additionally,  the  excavation  of  the  last 
abdominal  sternite  of  males  is  deeper  and  the 
margins  more  highly  elevated. 

The  coloration  of  parva  varies  from  brownish 
to  black  and  the  basal  oblique  dark  bands  of  the 
elytra  are  often  vague. 

Acyphoderes  suavis  Bates 

Acyphoderes  suavis  BATES,  1885:290,  pi.  20,  fig.  20. 

MALE:  Form  elongate;  integument  reddish; 
antennae  with  segments  6-8  usually  infuscated; 
elytra  pale  brownish  with  a  pale  triangular  basal 
area;  underside  variously  infuscated;  front  and 
middle  femora  basally  and  dorsally  dark,  hind 
femora  dark  at  middle;  hind  tibiae  dark  annulate 
on  apical  one-half.  Head  with  eyes  separated  on 
front  by  slightly  more  than  diameter  of  antennal 
scape.  Pronotum  longer  than  broad,  sides  lightly 
impressed  behind  middle;  apex  and  base  nar- 
rowly impressed;  dorsal  calluses  moderately  el- 
evated, opaque;  apex  and  base  with  a  band  of 
dense,  appressed  pubescence;  mesosternum 
with  epimeron  densely  pubescent;  metasternum 
with  a  densely  pubescent  patch  at  posterior  edge 
of  episternum.  Elytra  extending  to  third  abdom- 
inal segment;  lateral  margins  lightly  infuscated, 
densely  punctate;  disk  finely,  sparsely  punctate; 
apices  truncate,  as  broad  as  or  narrower  than 
antennal  scape.  Legs  with  anterior  and  inter- 
mediate femora  strongly  clavate,  posterior  pair 
gradually  clavate.  Abdomen  gradually  enlarged 


apically;  last  sternite  impressed  at  apex,  sides 
moderately  produced  apically.  Length,  13-24 
mm. 

FEMALE:  Form  similar.  Abdomen  more  bul- 
bous, last  sternite  not  impressed.  Length,  15-22 
mm. 

TYPE-LOCALITY:  Cordoba,  Mexico. 

FLOWER  RECORDS:  Buddleia,  Jatropha  (Mex- 
ico); Case  aria,  Baltimora  (Costa  Rica). 

This  species  may  be  readily  recognized  by  the 
color  and  pubescent  bands  of  the  pronotum. 

In  flight,  adults  greatly  resemble  and  behave 
like  those  of  Polistes  instabilis  Sauss. 

NEW  RECORDS:  See  Figure  6.  Known  from 
Mexico  to  Costa  Rica. 

Genus  Stenopseutes  Bates 

Stenopseutes  BATES,  1873:130;  1880:44. 

This  genus  may  be  recognized  by  the  slender 
antennae,  longitudinally  tricallose  pronotum,  re- 
duced prosternal  process,  and  slender,  feebly 
clavate  femora. 

TYPE-SPECIES:  Stenopseutes  aeger  Bates, 
1873  (monotypic). 

Two  species  are  known,  one  from  Mexico. 

Stenopseutes  sericinus  Bates 

Stenopseutes  sericinus  BATES,  1880:44. 

FEMALE:  Integument  testaceous,  antennae 
brownish.  Pronotum  densely  clothed  with  ap- 
pressed pubescence.  Elytra  vitreous  with  a  tri- 
angular patch  of  appressed  pubescence  at  base; 
margins  narrowly  dark.  Legs  yellowish,  femora 
feebly  clavate.  Length,  14  mm. 

TYPE-LOCALITY:  Mexico. 

No  new  material  of  this  species  has  been  seen. 
The  above  characterization  was  based  upon  a 
35-mm  color  transparency  of  a  specimen  in  the 
British  Museum  (Natural  History). 

Genus  Odontocera  Audinet-Serville 

Odontocera  AUDINET-SERVILLE,  1833:546;  WHITE  1855:187; 
THOMSON  1860:176;  1864:417;  LACORDAIRE  1869:503; 
BATES  1873:36;  1880:42;  LINSLEY  1963:165. 

The  primary  characteristic  of  this  genus  in 
Mexico  is  the  vitreous  disk  of  the  elytra.  Odon- 
tocera differs  from  Ommata  by  this  character 
and  also  by  the  less  elongate  pronotum.  Acy- 
phoderes differs  by  the  tricallused  pronotum  and 
usually  more  strongly  clavate  femora. 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


TYPE-SPECIES:  Odontocera  vitrea  Audinet- 
Serville  (Thomson  designation,  1864). 

Key  to  the  Mexican  species  of  Odontocera 

1.  Pronotum  margined  apically  and  basally 
by  bands  of  dense,  appressed  pubes- 
cence    2 

Pronotum  lacking  dense  pubescent 
bands,  disk  usually  with  a  longitudinal 
median  callus.  Length,  11-16  mm. 

Mexico  to  Nicaragua  clara 

2(1).  Elytra  more  than  3  times  longer  than 
basal  width,  erect  hairs  at  base  short; 
antennae  extending  beyond  middle  of 
elytra.  Length,  1 1-20  mm.  Southern 
Arizona  to  Chiapas  and  Veracruz  ____ 
aurocincta 

Elytra  less  than  3  times  longer  than  bas- 
al width,  erect  hairs  at  base  long;  an- 
tennae not  reaching  to  middle  of  ely- 
tra. Length,  12-13  mm.  Tamaulipas  to 
Veracruz fuscicornis 

Odontocera  fuscicornis  Bates 

Odontocera  fuscicornis  BATES,  1885:289. 

MALE:  Form  slender,  rather  short;  integument 
black;  antennae  reddish;  legs  brownish,  femora 
pale  at  bases;  elytra  yellowish,  vitreous,  nar- 
rowly margined  by  black,  apices  black,  base 
with  two  dark  humeral  vittae;  abdomen  with 
first  two  segments  partially  pale.  Head  with 
front  short,  eyes  contiguous;  antennae  short, 
segments  from  sixth  expanded  at  apices,  basal 
segments  with  a  few  long,  erect  hairs  beneath. 
Pronotum  moderately  coarsely,  subreticulately 
punctate;  base  deeply  impressed,  apical  and 
basal  pubescent  bands  silvery,  long,  erect  hairs 
numerous.  Elytra  less  than  3  times  longer  than 
basal  width:  punctures  on  vitreous  areas  sparse: 
pubescence  at  base  long,  erect.  Legs  with  fem- 
ora clavate;  posterior  tibiae  moderately  pubes- 
cent. Abdomen  with  last  sternite  impressed  for 
most  of  its  length,  sides  feebly  elevated.  Length, 
12-13  mm. 

FEMALE:  Form  and  coloration  similar.  Head 
with  eyes  well  separated  on  front.  Abdomen 
with  last  sternite  not  modified.  Length,  13  mm. 

TYPE-LOCALITY:  Cordoba,  Mexico. 

The  smaller  size,  shorter  elytra  and  antennae 
will  separate  this  species  from  aurocincta. 

NEW  RECORDS:  2  males.  1  female,  22  miles  [ca.  35  km]  N 
El  Limon,  Tamaulipas,  9  June  1951  (P.  D.  Hurd). 


Odontocera  aurocincta  Bates 

Odontocera  aurocincta  BATES,  1873:42. 

MALE:  Form  elongate,  slender,  integument 
black;  antennae  pale  with  basal  segments  often 
infuscated;  elytra  pale,  shining,  base  often 
black,  lateral  margins  black  to  beyond  middle; 
legs  black  or  with  only  clavate  portions  of  fem- 
ora black;  abdomen  with  segments  one  and  two 
mostly  pale.  Head  with  front  moderate,  eyes 
subcontiguous;  antennae  slender,  extending  a 
little  beyond  middle  of  elytra,  segments  from 
sixth  slightly  produced  apically.  Pronotum  long- 
er than  broad,  sides  rounded;  disk  coarsely,  sub- 
confluently  punctate;  apex  and  base  with  a  band 
of  dense,  silvery  or  golden,  appressed  pubes- 
cence, bands  extended  along  sides  and  united; 
long  erect  hairs  numerous;  mesosternum  dense- 
ly pubescent  on  epimeron;  pubescent  bands  on 
anterior  and  posterior  margins  of  metasternum 
and  on  episternum.  Elytra  more  than  3  times 
longer  than  basal  width;  margins  densely  punc- 
tate, pale  portion  of  disk  rather  sparsely,  sepa- 
rately punctate;  pubescence  sparse,  short,  erect 
over  basal  half.  Legs  slender,  femora  clavate. 
Abdomen  with  last  three  sternites  densely  punc- 
tate, last  sternite  feebly  impressed  over  most  of 
its  length,  sides  margined  but  not  elevated. 
Length,  1 1-19  mm. 

FEMALE:  Form  similar;  head  and  pronotum 
often  reddish;  antennae  usually  pale:  legs  pale, 
with  femoral  clavae  black;  abdomen  usually 
mostly  yellowish  or  reddish;  pubescence  golden. 
Head  with  eyes  widely  separated  on  front.  Ab- 
domen somewhat  inflated  toward  apex;  last  ster- 
nite rounded  at  apex.  Length,  12-20  mm. 

TYPE-LOCALITY:  Tehuantepec,  Mexico. 

This  species  is  easily  separated  by  the  elon- 
gate form  and  pubescent  bands  of  the  pronotum. 

Two  distinct  subspecies  can  be  recognized. 


Odontocera  aurocincta  aurocincta  Bates 

Odontocera  aurocincta  BATES,  1873:42:  1880:43. 

Odontocera  aurocincta  aurocincta',  LINSLEY  1961:163. 

Odontocera  aurocincta  var.  nigroapicalis  FISHER,  1947:52; 
LINSLEY  1961:164  (type-locality,  La  Gloria.  Cardel,  Vera- 
cruz). 

Males  with  pubescence  golden;  antennae  usu- 
ally all  pale;  elytra  without  basal  black  bands; 
legs  with  femoral  clubs  black.  Females  with 
head  and  pronotum  black. 

TYPE-LOCALITY:  Tehuantepec. 


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FIGURE  6.     Known  distribution  of  Acyphoderes  suavis  Bates  in  Mexico. 


RANGE:  State  of  Nayarit  to  Chiapas,  north 
to  Veracruz  (Fig.  7). 

FLIGHT  PERIOD:  June  to  August. 

Adults  have  been  taken  on  flowers  of  Acacia. 

Odontocera  aurocincta  arizonensis  Linsley 

Odontocera   aurocincta  arizonensis    LINSLEY,    1961:163; 
1963:165. 

Males  with  silvery  pubescence;  antennae  with 
basal  segments  infuscated;  legs  usually  all  black; 
elytra  black  basally.  Females  with  head  and 
pronotum  reddish. 

TYPE-LOCALITY:  Box  Canyon,  Santa  Rita 
Mts.,  Arizona. 

RANGE:  southern  Arizona  to  Sinaloa  (Fig.  7). 

FLIGHT  PERIOD:  July  to  September. 

Adults  frequent  the  flowers  of  Acacia,  Bud- 
dleia,  Croton,  and  Jatropha. 

Although  the  models  have  not  been  specifi- 
cally determined,  it  is  clear  that  the  two  sexes 
of  this  subspecies  mimic  different  insects.  Fe- 
males greatly  resemble  and  fly  like  vespids, 
while  males  appear  more  like  ichneumonids  in 
the  field. 


Odontocera  clara  Bates 

Odontocera  clara  BATES,  1873:38:  1874:222;  1880:43,  pi.  5, 

fig.  4. 
Odontocera  carinicollis  LINSLEY,  1934:348;  1935:84,  pi.  2,  fig. 

6  (type-locality:  Bejucos,  Temascaltepec,  Mexico).  NEW 

SYNONYMY. 

MALE:  Form  slender,  elongate;  integument 
black;  pronotum  often  reddish;  elytra  pale  vit- 
reous except  for  narrow  black  margins;  legs  oc- 
casionally pale;  antennae  often  with  outer  seg- 
ments pale  annulate.  Head  with  front  moderately 
long,  eyes  subcontiguous;  antennae  slender,  ex- 
tending beyond  middle  of  elytra,  segments  from 
sixth  slightly  produced  apically.  Pronotum  long- 
er than  broad,  sides  rounded;  disk  rather  finely, 
irregularly  punctate;  middle  usually  with  a  lon- 
gitudinal, slightly  elevated  callus  which  extends 
over  most  of  the  length:  pubescence  fine,  erect, 
with  appressed  pubescence  at  sides;  metaster- 
num  densely  pubescent.  Elytra  more  than  3l/2 
times  longer  than  basal  width;  base  and  black 
margins  densely  punctate,  pale  disk  sparsely 
punctate;  pubescence  sparse.  Legs  slender; 
femora  moderately  clavate.  Abdomen  densely 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


83 


FIGURE  7.     Known  distribution  of  Acyphode res  aurocincta  aurocincta  Bates  (closed  circles),  and  A.  aurocincta  arizonensis 
Linsley  (open  circles). 


pubescent;  last  sternite  shallowly  impressed  at 
apex,  sides  barely  elevated.  Length,  11-14  mm. 

FEMALE:  Form  similar.  Pronotum  a  little 
broader.  Abdomen  with  last  sternite  narrowly 
rounded  at  apex.  Length,  14-16  mm. 

TYPE-LOCALITY:  Chontales,  Nicaragua. 

This  species  varies  in  having  the  pronotum 
reddish  or  black.  The  elongate,  median  callus  of 
the  pronotum  characterizes  it. 

NEW  RECORDS:  1  male,  1  female,  6  miles  [ca.  10  km]  N  La 
Ventosa,  Oaxaca,  19  July  1963  (W.  A.  Foster);  1  female,  Su- 
chiapa,  Chiapas,  17  July  1957  (J.  A.  Chemsak);  1  female,  5.5 
miles  [ca.  8.9  km]  NW  Acala,  Chiapas,  23  June  1965  (Burke, 
Meyer,  Schaffner):  1  female,  30  miles  [ca.  48  km]  NE  Te- 
huantepec,  Oaxaca,  8  July  1955  (D.  Giuliani);  1  female,  4.4 
miles  [ca.  7.1  km]  E  Cuernavaca,  Morelos,  6-8  July  1974 
(Clark,  Murray,  Ashe,  Schaffner). 

Genus  Ommata  White 

Ommata  WHITE,  1855:194;  THOMSON  1864:166:  LACORDAIRE 
1869:502;  BATES  1870:319:  1873:26;  1880:42;  ZAJCIW 
1966:875;  1970:37. 

This  is  probably  the  most  difficult  genus  of 
Rhinotragini  to  define.  It  has  been  divided  into 


a  number  of  subgenera,  but  a  study  of  the  entire 
group  will  be  necessary  to  correctly  determine 
the  systematic  position  of  the  Mexican  Ommata. 
In  Mexico,  the  genus  may  be  recognized  by  the 
usually  long  antennae,  non-callused,  elongate 
pronotum  and  usually  non-vitreous  elytra. 

TYPE-SPECIES:  Ommata  elegans  White,  1855 
(monotypic). 

Three  species  are  presently  known  from  Mex- 
ico. 

Key  to  the  Mexican  species  of  Ommata 

1.  Pronotum  elongate,  cylindrical,  not  broad- 

ly impressed  at  base  and  apex;  elytra 
subglabrous,  with  fine  erect  hairs  arising 

from  each  puncture  2 

Pronotum  about  as  long  as  broad,  broadly 
impressed  at  base  and  apex;  elytra 
densely  clothed  with  appressed,  golden 
pubescence.  Length,  1 1  mm.  Veracruz 
to  Panama  sallaei 

2.  Integument,  especially  pronotum  and  un- 

derside, metallic  bluish  to  greenish;  pos- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  3 


terior  tibiae  densely  clothed  with  sub- 
erect,  black  setae.  Length,  11-13  mm. 

Veracruz  to  Oaxaca cyanea 

Integument  non-metallic,  abdomen  often 
reddish;  posterior  tibiae  sparsely  pubes- 
cent. Length,  7-10  mm.  Morelos  to  Gua- 
temala    championella 

Ommata  championella  Bates 

Ommata   (Eclipta)  championella    BATES,    1880:42;   ZAJCIW 

1970:38. 
Ommata  (Ommata)  rubriventris  LINSLEY,  1934:347;  1935:84; 

ZAJCIW   1966:91  (type-locality:  Tejupilco,  Temascal tepee, 

Mexico).  NEW  SYNONYMY. 

MALE:  Integument  black;  antennae  with  outer 
segments  basally  yellow  annulate;  elytra  with 
disk  longitudinally  pale  brownish;  femora  yel- 
lowish basally;  abdomen  often  reddish.  Head 
with  antennae  enlarged  from  seventh  segment. 
Pronotum  narrow,  shallowly  reticulate-punc- 
tate. Elytra  moderately  coarsely,  densely  punc- 
tate, punctures  becoming  denser  toward  apex. 
Abdomen  elongate;  last  sternite  impressed  at 
apex.  Length,  7-9  mm. 

FEMALE:  Form  more  robust.  Prothorax  often 
reddish.  Abdomen  broader,  reddish;  last  sternite 
narrowly  rounded  at  apex.  Length,  8-10  mm. 

TYPE-LOCALITY:  Calderas,  Guatemala. 

RANGE:  Morelos,  Mexico  to  Guatemala. 

The  small,  slender  form,  punctation  of  the 
pronotum,  and  paler  longitudinal  stripes  of  the 
elytra  will  separate  this  species  from  other 
known  Mexican  Ommata. 

NEW  RECORDS:  1  female,  7  miles  [ca.  1 1  km]  SW  Yautepec, 
Morelos,  2  July  1961,  3,500  ft  [ca.  1,067  m]  (Univ.  Kansas 
Mex.  Exped.);  1  male,  1  female,  3  miles  [ca.  5  km]  SE  La 
Trinitaria,  Chiapas,  18,  19  June  1965  (Burke,  Meyer,  Schaff- 
ner);  1  female,  Las  Margaritas,  Chiapas,  18  June  1965  (Burke, 
Meyer,  Schaffner);  1  female,  7  miles  [ca.  1 1  km]  SWTeopisca, 
Chiapas,  23  May  1969  (J.  M.  Campbell). 

Ommata  cyanea  Bates 

Ommata  cyanea  BATES,  1885:288. 
Ommata  (Eclipta)  cyanea;  ZAJCIW,  1970:38. 

MALE:  Integument  greenish  or  bluish  metallic; 
antennae  bluish,  yellow  annulate  from  fourth 
segment;  legs  bluish;  elytra  brownish  down  mid- 
dle. Head  with  antennae  as  long  as  elytra,  seg- 
ments from  sixth  apically  expanded.  Pronotum 
cylindrical;  coarsely  punctate  at  middle;  sparse- 
ly pubescent;  strongly  narrowed  toward  apex; 
apices  truncate.  Legs  with  posterior  femora 


elongate,  feebly  clavate.  Abdomen  with  last 
sternite  deeply  impressed.  Length,  11  mm. 

FEMALE:  Form  similar.  Antennae  black;  basal 
segments  densely  pubescent.  Abdomen  broader; 
last  sternite  narrowly  rounded  at  apex.  Length, 
12-13  mm. 

TYPE-LOCALITY:  Cordova,  Mexico. 

The  metallic  coloration  makes  this  species 
distinctive. 

NEW  RECORDS:  1  female,  5  miles  [ca.  8  km]  S  Lake  Cate- 
maco,  Veracruz,  6  July  1961  (D.  H.  Janzen);  1  female,  1  male, 
Temescal,  Oaxaca,  6  July  1965  (G.  Nelson  and  family). 

Ommata  sallaei  Bates 

Ommata  sallaei  BATES,  1885:289,  pi.  20,  fig.  15. 

FEMALE:  Integument  yellowish;  mandibles, 
eyes,  antennae  basally  and  at  apices  of  outer 
segments,  apical  and  basal  margins  and  rounded 
median  spot  on  pronotum,  elytra  except  for  tri- 
angular basal  area,  apices  and  dorsal  edges  of 
femora  and  bands  on  clubs  of  hind  femora,  and 
parts  of  underside  black.  Antennae  slender,  seg- 
ments feebly  produced  at  apices.  Eyes  widely 
separated  on  front.  Pronotum  coarsely  punctate, 
sparsely  pubescent.  Elytra  opaque,  densely 
punctate  and  pubescent,  apices  truncate.  Length, 
11  mm. 

TYPE-LOCALITY:  Cordova,  Veracruz. 

This  species  is  distinctive  among  Mexican 
Ommata  in  its  color  and  opaque  elytra.  It  is  also 
known  from  Cerro  Campana  in  Panama. 

ACKNOWLEDGMENTS 

These  studies  were  carried  out  in  conjunction 
with  a  National  Science  Foundation-sponsored 
study  on  North  American  Cerambycidae  through 
Grant  DEB  76-23849  A01.  The  authorities  of  the 
following  institutions  and  individuals  are  grate- 
fully acknowledged  for  the  loan  of  specimens: 
American  Museum  of  Natural  History,  New 
York;  California  Academy  of  Sciences,  San 
Francisco;  Canadian  National  Collection,  Otta- 
wa; Cornell  University,  Ithaca;  Essig  Museum 
of  Entomology,  Berkeley;  Field  Museum  of 
Natural  History,  Chicago;  Los  Angeles  County 
Museum  of  Natural  History;  Museum  of  Com- 
parative Zoology,  Cambridge:  Texas  A  &  M 
University,  College  Station;  United  States  Na- 
tional Museum  of  Natural  History,  Washington, 
D.C.;  University  of  Arizona,  Tucson;  Univer- 
sity of  California,  Davis;  University  of  Kansas, 


CHEMSAK  &  LINSLEY:  RHINOTRAGINI  OF  MEXICO 


85 


Lawrence;  E.  Giesbert,  D.  Marqua,  G.  Nelson, 
and  R.  Westcott. 

Celeste  Green  prepared  the  illustrations  and 
Kathleen  Sorenson  the  maps. 


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PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  4,  pp.  87-133,  70  figs.  December  22,  1979 


STUDIES  ON  THE  NEBRIINI  (COLEOPTERA:  CARABIDAE),  III.1 

NEW  NEARCTIC  NEBRIA  SPECIES  AND  SUBSPECIES, 
NOMENCLATURAL  NOTES,  AND  LECTOTYPE  DESIGNATIONS2 

By 
David  H.  Kavanaugh 

Department  of  Entomology,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  94118 


ABSTRACT:  The  purpose  of  this  paper,  third  of  a  series  on  the  nebriine  carabid  beetles,  is  to  upgrade  the 
status  of  nomenclature  for  Nearctic  members  of  genus  Nebria  in  preparation  for  subsequent  papers  on  classi- 
fication, phylogeny,  zoogeography,  and  natural  history  of  members  of  the  Nearctic  fauna.  Names  are  provided 
for  five  undescribed  species  [Nebria  carri  (type-locality — Dollarhide  Summit,  Idaho),  V.  darlingtoni  (type-lo- 
cality— South  Fork  American  River,  California),  N.  gouleti  (type-locality — Rattlesnake  Creek,  Washington),  N. 
lituyae  (type-locality — Mount  Blunt,  Lituya  Bay,  Alaska),  and  N.  navajo  (type-locality — 19  miles  SW  of  Kayenta, 
Arizona)].  Twenty-three  subspecies  are  also  described  as  new  [Nebria  acuta  quileute,  N.  arkansana  edwardsi, 
N.  arkansana  oowah,  N.  arkansana  uinta,  N.  fragilis  teewinot,  N.  gebleri  cascadensis,  N.  gebleri  fragariae,  N. 
gebleri  siskiyouensis,  N.  gyllenhali  lassenensis,  N.  gyllenhali  lindrothi,  N.  kincaidi  balli,  N.  lacustris  bellorum,  N. 
meanyi  lamarckensis,  N.  meanyi  sylvatica,  N.  nivalis  gaspesiana,  N.  obliqua  chuskae,  N.  sahlbergii  modoc,  N. 
sahlbergii  triad,  N.  schwarzi  beverlianna,  N.  spatulata  sierrae,  N.  trifaria  utahensis,  N.  vandykei  wyeast,  and  N. 
zioni  oasis].  Diagnosis  and  comment  on  geographical  distribution  are  provided  for  each  new  taxon.  Lectotypes 
are  designated  for  Carabus  gyllenhali  Schonherr,  C.  nivalis  Paykull,  Helobia  castanipes  Kirby,  Nebria  bifaria 
Mannerheim,  N.  eschscholtzii  Menetries,  N.  gregaria  Fischer  von  Waldheim,  N.  hudsonica  LeConte,  N.  ingens 
Horn,  A7,  livida  LeConte,  N.  mannerheimii  Fischer  von  Waldheim,  A',  metallica  Fischer  von  Waldheim,  N.  mollis 
Motschulsky,  N.  moesta  LeConte,  N.  obliqua  LeConte,  N.  sahlbergii  Fischer  von  Waldheim,  N.  suturalis 
LeConte,  N.  trifaria  LeConte,  N.  vandykei  Banninger,  N.  violacea  Motschulsky,  and  N.  viridis  Horn.  Nebria 
longula  LeConte  and  N.  obtusa  LeConte  are  recognized  as  junior  synonyms  of  A',  suturalis  LeConte  and  N. 
obliqua  LeConte,  respectively.  Nebria  rathvoni  LeConte  is  reduced  in  status  to  a  subspecies  of  W.  gebleri  Dejean. 
Emendations  or  restrictions  of  type-localities  are  provided  for  Nebria  catenata  Casey,  N.  eschscholtzii  Menetries, 
N.  expansa  Casey,  N.  hudsonica  LeConte,  N.  incerta  Casey,  N.  ingens  Horn,  N.  longula  LeConte,  N.  obliqua 
LeConte,  N.  oregona  Casey,  N.  ovipennis  LeConte,  and  N.  tenuipes  Casey.  Additional  notes  on  type-specimens 
and  nomenclature  are  provided  for  other  species  group  names. 


INTRODUCTION 


1  Papers  which  serve  as  contributions  I  and  II  in  this  series  My  interest  in  beetles  of  the  genus   Nebria 
are,  respectively,  Kavanaugh  (1971)  and  Kavanaugh  and  Mar-  Latreille  began  in  1967.  While  collecting  Carabid 

tmko  (1972).  beetles  in  the  Rocky  Mountains  of  Colorado,  I 

2  Data  presented  here  have  been  extracted  from  the  author's  .     .                                        r          ,    , 

....,„.         .  _.      '  t.  concentrated  on  my  own  preferred  haunts — 

unpublished  Doctoral  Dissertation,  submitted  to  the  Univer-  * 

sity  of  Alberta,  Edmonton,  September  1978  (Kavanaugh  namely,  riparian,  nival,  and  pen-nival  habitats. 

1978).  This  specialization  provided  frequent  encoun- 

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ters  with  members  of  various  Nebria  species 
because,  in  these  habitats,  few  insect  groups  ap- 
proach Nebria  in  their  conspicuous  abundance. 
With  the  aid  of  Lindroth's  (1961)  treatment  of 
the  genus  for  Canada  and  Alaska,  I  was  able  to 
identify  most  of  my  material.  It  soon  became 
evident  that  Nebria  species  formed  a  biologi- 
cally important  and  interesting  element  of  the 
montane  fauna.  Species  diversity  and  population 
densities  in  suitable  microhabitats  were  impres- 
sively high,  and  I  began  to  recognize  patterns  of 
altitudinal  zonation  among  the  resident  species. 
Observations  made  on  collecting  trips  to  mon- 
tane areas  in  Utah  and  California  suggested  that 
similar  patterns  of  Nebria  species  diversity  and 
habitat  distribution  existed  in  these  and  other 
areas  in  western  North  America. 

The  opportunity  to  begin  serious  study  of 
Nearctic  Nebria  appeared  in  July  1970,  when  I 
enrolled  at  the  University  of  Alberta,  Edmon- 
ton. Although  Nearctic  Nebria  species  were  at 
that  time  already  among  the  taxonomically  bet- 
ter-known carabid  groups,  I  hoped  to  build  on 
my  previous  interest  and  experience  through  ad- 
ditional studies  which  seemed  both  feasible  and 
potentially  rewarding  scientifically.  My  plan  was 
to  obtain  data  with  which  to  further  define  the 
structural  and  distributional  limits  of  Nearctic 
species  and  explore  the  evolutionary  relation- 
ships among  them.  Initial  goals  were  the  for- 
mulation of  an  improved  infrageneric  classifi- 
cation of  the  New  World  species  and  generation 
of  hypotheses  concerning  the  historical  devel- 
opment of  diversity  and  distributional  patterns 
observed  in  the  fauna. 

Nebria,  in  the  broadest  sense,  comprises  a 
group  of  over  500  nominal  taxa  which,  in  aggre- 
gate, demonstrate  Holarctic  distribution.  The 
Palaearctic  component  of  the  genus  is  more  di- 
verse, both  in  species  and  major  lineages,  than 
the  Nearctic.  Consequently,  all  hypotheses  on 
relationships  among  Nearctic  species  and  on 
historical  development  of  the  Nearctic  fauna 
would  be  highly  speculative  in  the  absence  of  a 
working  knowledge  of  the  Palaearctic  fauna. 
Fortunately,  as  work  toward  the  initial  project 
goals  progressed,  new  sources  of  taxonomic 
data  became  available  as  by-products  of  field 
work  and  efforts  to  obtain  comparative  material 
on  loan.  These  additional  resources  have  per- 
mitted expansion  of  the  project  to  include  cov- 
erage of  the  Palaearctic  Nebria  fauna,  compar- 


isons with  members  of  other  nebriine  genera  and 
related  carabid  tribes,  and  the  use  of  additional 
character  systems  in  study  of  the  Nearctic  Ne- 
bria fauna.  These  other  systems  include  external 
structure  of  immature  stage  individuals,  life  his- 
tory features  (especially  life  cycle  timing),  hab- 
itat preferences,  and  adult  internal  structure. 

Various  facets  of  the  expanded  project  have 
progressed  at  different  rates,  and  some  of  these 
are  yet  incomplete  or  barely  begun.  This  paper 
represents  a  first  report  on  the  Nearctic  Nebria 
fauna,  specifically  on  nomenclature  relating  to 
same.  My  purpose  is  twofold.  First,  I  provide 
names  for  five  species  and  23  subspecies  not 
previously  described.  These  names  are  needed 
immediately  for  use  in  several  other  manuscripts 
and  by  several  other  workers.  To  this  end,  data 
and  discussion  presented  for  each  name  are  lim- 
ited to  little  more  than  the  minimum  required  by 
the  International  Code  of  Zoological  Nomencla- 
ture for  availability.  My  second  purpose  is  to 
provide  additional  notes  on  nomenclature  for 
previously  described  Nearctic  species,  including 
emendations  and/or  restrictions  of  type-locali- 
ties, notes  on  type-specimens,  and  designations 
of  lectotypes. 

In  a  subsequent  paper  (manuscript  in  prepa- 
ration) I  will  present  additional  information  on 
Nearctic  Nebria  taxa,  including  those  presented 
here  as  new.  Each  taxon  will  be  more  fully  de- 
scribed and  illustrated.  A  classification  and  key 
for  identification  of  adults  will  be  provided,  as 
will  results  of  comparative  studies  on  structure, 
distribution,  and  natural  history.  A  review  of  the 
known  fossil  record  of  Nearctic  Nebria  will  also 
be  included. 

MATERIALS 

This  study  is  based  on  examination  of  over 
66,000  Nearctic  and  5,800  Palaearctic  adult  Ne- 
bria specimens.  Approximately  35,000  speci- 
mens were  borrowed  from  various  institutional 
and  private  collections  in  North  America,  Eu- 
rope, and  Asia.  Following  is  a  list  of  abbrevia- 
tions used  in  the  text  which  refer  to  various  col- 
lections from  which  specimens  were  received. 
Most,  but  not  all,  correspond  to  abbreviations 
proposed  by  Arnett  and  Samuelson  (1969). 
Where  appropriate,  names  of  curators  or  assis- 
tants who  sent  specimens  are  also  included. 

AHab — A.  Habu,  National  Institute  of  Agricultural  Sciences, 
Tokyo,  Japan. 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


ALar — A.  Larochelle,  College  Bourget,  Rigaud,  Quebec  JOP 
IPO. 

AMor — A.  Morgan  and  A.  V.  Morgan,  University  of  Water- 
loo, Waterloo,  Ontario  N2L  3G1. 

AMNH — American  Museum  of  Natural  History,  New  York, 
New  York  10024;  L.  H.  Herman,  Jr. 

ANSP— Academy  of  Natural  Sciences,  Philadelphia,  Penn- 
sylvania 19103;  W.  W.  Moss. 

BFCa— B.  F.  Carr,  24  Dalrymple  Green  NW,  Calgary,  Al- 
berta T3A  1Y2. 

BMNH — British  Museum  (Natural  History),  London,  En- 
gland; P.  M.  Hammond,  H.  K.  Kenward,  M.  E.  Bacchus. 

BRot— B.  Rotger,  CR.,  Immaculate  Heart  of  Mary  Church, 
P.O.  Box  451,  Pagosa  Springs,  Colorado  81147. 

CArm— C.  Armin,  191  West  Palm  Avenue,  Reedley,  Califor- 
nia 93654. 

CAS — California  Academy  of  Sciences,  San  Francisco,  Cali- 
fornia 941 18;  H.  B.  Leech. 

CDA — California  State  Department  of  Food  and  Agriculture, 
Sacramento,  California  95814;  F.  G.  Andrews. 

CNC— Canadian  National  Collection  of  Insects,  Biosystema- 
tics  Research  Institute,  Ottawa,  Ontario  K1A  OC6;  R.  de 
Ruette,  A.  Smetana. 

CSU— Colorado  State  University,  Fort  Collins,  Colorado 
8052 1;T.  O.  Thatcher. 

CUB— University  of  Colorado,  Boulder,  Colorado  80302;  H. 
Rodeck. 

CUIC— Cornell  University,  Ithaca,  New  York  14850;  L.  L. 
Pechuman. 

DBUM—  Universite  de  Montreal,  Montreal,  Quebec  H3C  3J7; 
M.  Coulloudon. 

DENH — University  of  New  Hampshire,  Durham,  New 
Hampshire  03824;  W.  J.  Morse. 

DEUN— University  of  Nebraska,  Lincoln,  Nebraska  68503; 
B.  C.  Ratcliffe. 

DHKa — D.  H.  Kavanaugh,  California  Academy  of  Sciences, 
San  Francisco,  California  94118. 

DJLa — D.  J.  Larson,  Department  of  Biology,  Memorial  Uni- 
versity of  Newfoundland,  St.  John's,  Newfoundland  A  1C 
5S7. 

DMan — D.  Mann,  University  of  Washington,  Seattle,  Wash- 
ington 98195. 

DRWh— D.  R.  Whitehead,  Systematic  Entomology  Labora- 
tory, U.S.D.A.,  %  United  States  National  Museum,  Wash- 
ington D.C.  20560. 

DZEC — Montana  State  University,  Bozeman,  Montana 
59715;  N.  L.  Anderson. 

EAMa — E.  A.  Martinko,  University  of  Kansas,  Lawrence, 
Kansas  66045. 

EDNC — North  Carolina  Department  of  Agriculture,  Raleigh, 
North  Carolina  27602;  J.  F.  Greene. 

EMUS— Utah  State  University,  Logan,  Utah  84332;  W.  J. 
Hanson. 

ETHZ — Eidgenossische  Technische  Hochschule  Entomolo- 
gisches  Institut,  Zurich,  Switzerland;  W.  Sauter. 

FMNH — Field  Museum  of  Natural  History,  Chicago,  Illinois 
60605;  H.  Dybas. 

GASh— G.  A.  Shook,  1209  West  Hays,  Boise,  Idaho  83702. 

GRNo — G.  R.  Noonan,  Milwaukee  Public  Museum,  Milwau- 
kee, Wisconsin  53233. 

HGou — H.  Goulet,  Biosystematics  Research  Institute,  Otta- 
wa, Ontario  K1A  OC6. 


HoKn— R.  L.  Hoffman  and  L.  Knight,  Radford  College,  Rad- 
ford,  Virginia  24141. 

HNHM — Hungarian  Natural  History  Museum,  Budapest, 
Hungary;  S.  Horvatovitch. 

ICCM— Carnegie  Museum,  Pittsburg,  Pennsylvania  15213;  G. 
E.  Wallace. 

INHS — Illinois  Natural  History  Survey,  Urbana,  Illinois 
61803;  M.  W.  Sanderson. 

ILar — I.  La  Rivers,  University  of  Nevada,  Reno,  Nevada 
89507. 

ISUI— Iowa  State  University,  Ames,  Iowa  50010;  R.  Miller. 

IUIC — Indiana  University,  Bloomington,  Indiana  47401;  F. 
N.  Young. 

JBel— J.  Belicek,  9345-98A  Street,  Edmonton,  Alberta  T6E 
3N2. 

JKus — J.  Kuster,  University  of  Alberta,  Edmonton,  Alberta 
T6G  2E3. 

JNeg — J.  Negre,  9  Boulevard  de  Lesseps,  Versailles,  France. 

JSch— J.  Schuh,  4039  Shasta  Way,  Klamath  Falls,  Oregon 
97601. 

JSpe — J.  Spence,  University  of  Alberta,  Edmonton,  Alberta 
T6G  2E3. 

JVMa — J.  V.  Matthews,  Jr.,  Geological  Survey  of  Canada, 
Ottawa,  Ontario  K1A  OE8. 

KMFe — K.  M.  Fender,  911  Ashwood  Avenue,  McMinnville, 
Oregon  97128. 

KSUC— Kansas  State  University,  Manhattan,  Kansas  66502; 
H.  D.  Blocker. 

KUSM — University  of  Kansas,  Snow  Museum,  Lawrence, 
Kansas  66045;  G.  Byers. 

LACM — Los  Angeles  County  Museum  of  Natural  History, 
Los  Angeles,  California  90007;  C.  L.  Hogue. 

LEMC — Macdonald  College,  Lyman  Entomological  Museum, 
Ste.  Anne  de  Bellevue,  Quebec  J7V  3N8. 

LRus— L.  Russell,  828  NW  27th  Street,  Corvallis,  Oregon 
97330. 

MCZ — Museum  of  Comparative  Zoology,  Harvard  Universi- 
ty, Cambridge,  Massachusetts  02138;  J.  F.  Lawrence. 

MGF — Museum  G.  Frey,  Entomologisches  Institut,  Tutzing 
bei  Miinchen,  West  Germany;  M.  Wiirmli. 

MHNG — Museum  d'Histoire  Naturelle,  Geneva,  Switzer- 
land; C.  Besuchet. 

MNHP — Museum  National  d'Histoire  Naturelle,  Paris, 
France;  A.  Bons,  H.  Perrin. 

MLLu — M.  L.  Luff.  University  of  Newcastle,  Newcastle- 
upon-Tyne,  England. 

MSU — Michigan  State  University,  East  Lansing,  Michigan 
48823;  R.  L.  Fischer. 

NMB — Naturhistorisches  Museum  Basel,  Basel,  Switzerland; 
W.  Wittmer. 

NMDo — N.  M.  Downie,  505  Lingle  Terrace,  Lafayette,  In- 
diana 47901. 

NMPC — National  Museum  in  Prague,  Prague,  Czechoslova- 
kia; Z.  Mlynar. 

NRSS — Naturhistoriska  Riksmuseet,  Stockholm,  Sweden;  T. 
Nyholm. 

NSDA — Nevada  State  Department  of  Agriculture,  Reno,  Ne- 
vada 89504;  R.  C.  Bechtel. 

ODA — Oregon  State  Department  of  Agriculture,  Salem,  Or- 
egon 97310;  R.  L.  Westcott. 

OSEC— Oklahoma  State  University,  Stillwater,  Oklahoma 
74074;  W.  A.  Drew. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


OSUC— Ohio  State  University,  Columbus,  Ohio  43210;  C.  A. 
Triplehorn. 

OSUO — Oregon  State  University,  Corvallis,  Oregon  97331;  P. 
Oman. 

PADA — Pennsylvania  State  Department  of  Agriculture,  Har- 
risburg,  Pennsylvania  17120;  T.  J.  Henry. 

PMCh— P.  M.  Choate,  Jr.,  University  of  Florida,  Gainesville, 
Florida  32601. 

PMNH — Peabody  Museum  of  Natural  History,  Yale  Univer- 
sity, New  Haven,  Connecticut  06520;  C.  L.  Remington,  K. 
W.  Brown. 

PSMi— P.  S.  Miliotis,  Depot  Street,  Dunstable,  Massachu- 
setts 01827. 

PSUC — Pennsylvania  State  University,  University  Park, 
Pennsylvania  16802;  K.  C.  Kim. 

PUCA— Pacific  Union  College,  Angwin,  California  94508;  L. 
E.  Eighme. 

PURC— Purdue  University,  Lafayette,  Indiana  47907;  A.  Pro- 
vonsha. 

RCGr— R.  C.  Graves,  627  Crestview,  Bowling  Green,  Ohio 
43402. 

RCra — R.  Crawford,  University  of  Washington,  Seattle, 
Washington  98195. 

RDav — R.  Davidson,  University  of  Vermont,  Burlington, 
Vermont  05401. 

RFre — R.  Freitag,  Lake  head  University,  Thunder  Bay,  On- 
tario P7B  5E1. 

ROM— Royal  Ontario  Museum,  Toronto,  Ontario  M5S  2C6; 
G.  B.  Wiggins. 

RPPa— R.  P.  Papp,  Bernice  P.  Bishop  Museum,  Honolulu, 
Hawaii  96818. 

RTBe— R.  T.  Bell,  University  of  Vermont,  Burlington,  Ver- 
mont 05401. 

SFVS — California  State  University,  Northridge,  California 
91324;  P.  F.  Bellinger. 

SDSU— South  Dakota  State  University,  Brookings,  South 
Dakota  57006;  E.  U.  Balsbaugh,  Jr. 

SJSU— San  Jose  State  University,  San  Jose,  California  951 14; 
J.  G.  Edwards. 

TCBa— T.  C.  Barr,  Jr.,  University  of  Kentucky,  Lexington, 
Kentucky  40506. 

TNak — T.  Nakane,  National  Science  Museum,  Tokyo,  Japan. 

TMZM — Troms0  Museum,  Tromsp,  Norway;  J.  H.  Ander- 
sen. 

UAFA — University  of  Arkansas,  Fayetteville,  Arkansas 
72701;  R.  T.  Allen. 

UASM — University  of  Alberta,  Strickland  Museum,  Edmon- 
ton, Alberta  T6G  2E3;  G.  E.  Ball. 

UBC — University  of  British  Columbia,  Spencer  Museum, 
Vancouver,  British  Columbia  V6T  1W5;  G.  G.  E.  Scudder. 

UCB— University  of  California,  Essig  Museum  of  Entomol- 
ogy, Berkeley,  California  94720;  J.  A.  Chemsak. 

UCD— University  of  California,  Davis,  California  95616;  R. 
O.  Schuster. 

UCR— University  of  California,  Riverside,  California  92502; 
S.  Frommer. 

UIMI— University  of  Idaho,  Moscow,  Idaho  83843;  W.  F. 
Barr. 

UMHF — Universitetets  Zoologiska  Museum  Entomologiska 
Avdelningen,  Helsingfors,  Finland;  H.  Silfverberg. 

UMMZ — University  of  Michigan,  Ann  Arbor,  Michigan 
48104;  R.  D.  Alexander. 

UMRM — University  of  Missouri,  Columbia,  Missouri  65201; 
W.  R.  Enns. 


UMTF— University  of  Turku,  Turku  (Abo),  Finland;  H.  Hip- 
pa. 

USNM — United  States  National  Museum  of  Natural  History, 
Smithsonian  Institution,  Washington,  D.C.  20560;  T.  L. 
Erwin. 

UWBM— University  of  Washington,  Burke  Museum,  Seattle, 
Washington  98105;  M.  H.  Hatch,  S.  Rohwer. 

UWEM — University  of  Wisconsin,  Madison,  Wisconsin 
53706;  L.  J.  Bayer. 

UWLW — University  of  Wyoming,  Laramie,  Wyoming  83070; 
R.  J.  Lavigne. 

VMKi— V.  M.  Kirk,  Northern  Grain  Insects  Research  Lab- 
oratories, Brookings,  South  Dakota  57006. 

WSU— Washington  State  University,  Pullman,  Washington 
99163;  W.  J.  Turner. 

ZILR— Zoological  Institute,  Academy  of  Science,  Leningrad, 
U.S.S.R.;  O.  L.  Kryzhanovskij,  V.  G.  Shilenkov. 

ZMKD — Universitetets  Zoologiske  Museum,  K0benhavn, 
Denmark;  O.  Martin. 

ZMLS — Zoological  Institute,  University  of  Lund,  Lund,  Swe- 
den; C.  H.  Lindroth. 

ZMUM — Zoological  Museum,  Moscow  University,  Moscow, 
U.S.S.R.;  S.  Keleinikova. 

ZSBS— Zoologische  Sammlung  des  Bayerischen  Staates, 
Miinchen  19,  West  Germany;  G.  Scherer. 

I  have  been  able  to  study  the  type-specimens 
of  all  85  previously  described  Nearctic  nominal 
taxa  through  the  cooperation  of  many  of  the  cu- 
rators listed  above.  The  following  three  minor 
nomenclatural  problems  must  remain  unsolved, 
but  none,  I  think,  will  require  further  attention. 
Nebria  melsheimeri  Sturm  (1826:173)  and  Ne- 
bria  eschscholtzii  Sturm  (1826:173)  (not  of  Me- 
netries)  must  be  considered  nomina  nuda  be- 
cause these  names  appear  only  in  Sturm's 
catalog  to  his  collection,  unaccompanied  by 
either  description  or  statement  of  locality;  and 
no  type-specimens  exist.  The  problem  with  and 
current  status  of  the  name  Nebria  elias  Mot- 
schulsky  is  discussed  with  a  treatment  of  Nebria 
gyllenhali  (Schonherr)  below. 

Where  necessary,  lectotypes  have  been  cho- 
sen and  so  labelled,  and  their  formal  designa- 
tions appear  here.  I  have  also  labelled  holotypes 
for  previously  described  species  names  where 
prior  labels  were  unclear  or  lacking.  Holotypes 
designated  for  taxa  described  here  as  new  (with 
the  exception  of  the  holotype  of  N.  trifaria  utah- 
ensis  n.  ssp.)  have  been  deposited  in  the  type 
collection  of  the  California  Academy  of  Sci- 
ences. 


METHODS 

A  detailed  discussion  of  preparative  and  pro- 
cedural methods  used  in  this  project  will  be  in- 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


91 


eluded  in  a  subsequent  paper  (Kavanaugh, 
manuscript  in  preparation).  I  present  here  com- 
ment on  only  those  methods  which  relate  spe- 
cifically to  data  and  results  presented  in  this  pa- 
per. 

DISSECTING  TECHNIQUES. — Characters  used 
in  diagnoses  of  many  taxa  named  here  include 
form  of  the  hindwings  and  genitalia  of  adults. 
Successful  examination  of  these  structures  re- 
quired some  dissection.  Specimens  were  pre- 
pared for  dissection  by  soaking  them  for  5  or  10 
minutes  in  boiling  water.  Addition  of  a  very 
small  amount  of  liquid  detergent  hastened  the 
relaxation  process.  Specimens  collected  in  ethyl 
acetate  fumes  were  relaxed  easily,  but  material 
collected  in  potassium  cyanide  fumes  or  in  ethyl 
alcohol  required  a  brief  (two  to  five  minute) 
treatment  in  warm  potassium  hydroxide  (10% 
solution). 

A  general  impression  of  the  size  (but  not 
shape)  of  the  hindwings  was  obtained  by  simply 
lifting  the  left  elytron  to  expose  the  wing.  For 
a  study  of  venation  and  wing  shape,  the  left 
hindwing  was  torn  free  at  the  wing  base  using 
fine  forceps,  then  spread  and  mounted  in  alcohol 
on  a  slide.  Permanent  mounts  (in  Euparal  me- 
dium) were  made  for  representatives  of  all  taxa 
studied;  however,  most  hindwings  were  later 
dried,  glued  to  cards,  and  pinned  with  their  re- 
spective specimens. 

The  apex  and  shaft  of  the  median  lobe  and 
parameres  in  male  specimens  or  the  coxostyli  in 
female  specimens  were  quickly  examined  in  re- 
laxed specimens  by  simple  eversion  of  these 
structures  using  fine  forceps.  For  study  of  the 
base  of  the  median  lobe  in  males  or  the  sper- 
matheca,  bursa  copulatrix,  and  other  internal 
structures  in  females,  the  genitalia  were  extract- 
ed as  a  unit  by  tearing  the  membranous  connec- 
tions between  eighth  tergum  and  sternum  and 
the  "ring  sclerite"  (in  males)  or  the  valvifers, 
paraprocts,  and  proctiger  (in  females).  Genitalia 
were  next  treated  for  5  to  10  minutes  in  hot 
(10%)  potassium  hydroxide  solution  to  remove 
nonsclerotized  tissue,  rinsed  in  water,  further 
dissected  as  needed,  then  transferred  to  and  ex- 
amined in  glycerine.  After  genitalic  specimens 
were  studied  and  drawn,  they  were  placed  in 
polyethylene  microvials  in  a  drop  of  glycerine 
and  pinned  with  their  respective  specimens. 

SEX  DETERMINATION. — Determination  of  the 
sex  of  individuals  was  made  simply  by  reference 


to  form  of  the  front  tarsi.  Adult  males  have  the 
basal  three  or  four  tarsomeres  dilated,  laterally 
expanded,  and  have  pads  of  adhesive  setae  on 
ventral  surfaces  of  the  basal  two  or  three  tar- 
someres. Females  have  all  tarsomeres  slender 
and  without  ventral  pads  of  adhesive  setae. 

MEASUREMENTS. — The  only  mensural  char- 
acter used  in  this  study  is  "standardized  body 
length."  This  expression  refers  to  the  sum  of 
three  measurements  (Fig.  1A):  length  of  head, 
measured  along  midline  from  apical  margin  of 
clypeus  to  a  point  opposite  posterior  margin  of 
eye;  length  of  pronotum,  measured  along  mid- 
line  from  apical  to  basal  margin;  and  length  of 
elytron,  measured  along  midline  from  apex  of 
scutellum  to  a  point  opposite  elytral  apex.  While 
this  measure  of  relative  size  avoids  error  due  to 
variation  in  extension  or  retraction  of  body  parts 
among  specimens,  it  consistently  underesti- 
mates the  apparent  total  body  length  of  speci- 
mens (such  as  could  be  measured  from  the  most 
anterior  to  the  most  posterior  point)  by  from  12 
to  18  percent. 

Range  in  standardized  body  length  in  my  sam- 
ple for  each  taxon  was  established  by  visual  se- 
lection of  smallest  and  largest  specimens  for 
each  sex.  All  measurements  were  made  with  the 
aid  of  a  Leitz  stereoscopic  dissecting  micro- 
scope at  a  magnification  of  16  diameters,  using 
a  calibrated  ocular  grid  with  a  scale  interval  of 
0.1  mm. 

ILLUSTRATIONS  AND  MAPS. — Line  drawings 
illustrating  structural  characters  were  made  with 
the  aid  of  an  ocular  grid  mounted  in  a  Leitz 
stereoscopic  dissecting  microscope.  Unless  oth- 
erwise noted,  scale  lines  which  accompany  il- 
lustrations equal  1.0  mm.  All  drawings  of 
homologous  structures  are  consistent  in  scale 
and  aspect  to  facilitate  comparisons.  In  some 
drawings,  uniform  sparse  stippling  is  used  to  in- 
dicate membranous  areas.  Where  only  part  of  a 
structure  is  illustrated  (such  as  the  basal  region 
of  an  elytron),  a  wavy  line  serves  as  the  artificial 
margin  of  the  drawing. 

Maps  are  presented  to  illustrate  known  distri- 
butions of  taxa.  Symbols  used  denote  approxi- 
mate locations  of  samples  (see  taxon  descrip- 
tions for  respective  lists  of  sample  localities). 
(Written  permission  has  been  obtained  for  use 
of  Goode  Base  Map  No.  202  [or  parts  thereof] 
[copyright  by  the  University  of  Chicago,  De- 
partment of  Geography].) 


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FORMAT  FOR  PRESENTATION  OF  NEW  TAXA. — 
Included  for  each  new  name  are:  a  synonymy 
(i.e.,  list  of  names  which  have  been  misapplied 
in  the  literature  to  members  of  this  taxon,  in- 
cluding all  known  literature  records);  designa- 
tions and  listings  of  type-specimens  and  their 
places  of  deposition;  designation  of  type-locali- 
ty; the  diagnostic  combination  (i.e.,  those  char- 
acter states  which,  in  combination,  distinguish 
members  of  the  taxon  from  members  of  other 
taxa);  derivation  of  the  taxon  name;  a  brief  syn- 
opsis of  geographical  distribution;  and  a  list  of 
localities  (including  months  of  collection  and 
place  of  deposition)  for  paratypes. 

Several  comments  are  required  on  the  format 
and  data  content  of  the  locality  lists.  For  con- 
venience, all  data  (including  countries,  states  or 
provinces,  counties,  and  localities)  are  arranged 
alphabetically.  All  records  are  from  specimens 
I  have  personally  studied.  Records  solely  from 
the  literature  are  not  included  here.  The  geo- 
graphical subdivisions  "County"  or  "Comte" 
are  used  where  possible  as  an  aid  in  organizing 
the  data.  Provinces  and  states  for  which  these 
subdivisions  are  not  used  include:  Alberta,  Brit- 
ish Columbia,  Manitoba,  Saskatchewan,  Yukon 
Territory,  and  Alaska.  All  national  parks  are 
treated  as  counties;  and  all  localities  within  park 
boundaries  are  listed  under  the  park  rather  than 
under  their  appropriate  counties.  Because  alti- 
tude data  are  often  important  in  defining  the  hab- 
itat ranges  of  Nebria  taxa,  these  data  are  re- 
corded here  where  known.  No  attempt  has  been 
made,  however,  to  present  altitude  data  not  ac- 
tually recorded  on  the  specimen  labels.  Where 
necessary,  altitude  values  have  been  converted 
to  their  metric  equivalents. 

Only  the  months  in  which  specimens  were  ac- 
tually collected  at  a  given  locality  have  been  not- 
ed here.  This  level  of  precision  adequately  de- 
scribes the  temporal  distributions  of  most 
samples.  The  addition  of  year  and  day  data 
would  have  expanded  the  size  of  this  paper 
greatly  but  contributed  little.  I  have  also  omitted 
the  names  of  collectors  in  order  to  conserve 
space.  Collections  in  which  specimens  are  de- 
posited are  noted  for  each  locality,  but  holdings 
for  each  collection  are  not  itemized. 

Specimens  with  illegible  or  essentially  mean- 
ingless locality  labels  are  grouped  with  speci- 
mens bearing  no  locality  labels  under  the  head- 
ing "SPECIMENS  WITHOUT  LOCALITY  DATA." 
Specimens  listed  under  the  heading  "DOUBTFUL 


RECORDS"  are  of  two  types:  (1)  those  labelled 
as  from  localities  outside  the  known  and  prob- 
able ranges  of  the  taxon;  and  (2)  those  labelled 
as  from  localities  within  the  known  geographical 
range  of  a  taxon  but  from  areas  where  I  am  cer- 
tain no  suitable  habitat  for  the  beetles  exists. 
For  some  specimens  in  category  (2),  however, 
where  appropriate  habitat  can  be  found  within 
reasonable  proximity  of  the  stated  locality,  I 
have  chosen  to  add  the  word  "area"  after  the 
entry.  Although  imprecise,  this  convention  ac- 
commodates the  common  practice  among  some 
collectors  of  simply  recording  the  nearest  city 
or  town  on  specimen  labels  rather  than  more 
precise  locality  data. 

CRITERIA  FOR  RANKING  TAXA. — All  available 
evidence  supports  the  universality  of  sexual  re- 
production among  nebriine  species.  Because  ap- 
parently all  Nebria  species  are  bisexual,  I  ac- 
cept as  appropriate  Mayr's  (1969:26)  proposed 
definition  of  the  biological  species  (as  amended 
by  Whitehead  1972:139),  namely,  that  species 
are  populations  or  groups  of  populations  through 
which  gene  flow  actually  or  potentially  exists, 
but  which  are  reproductively  [intrinsically]  iso- 
lated from  all  other  such  populations  [or  groups 
of  populations].  As  noted  by  Whitehead 
(1972:139)  and  Larson  (1975:251),  the  species  so 
defined  is  a  category  with  a  nonarbitrary,  objec- 
tive basis;  but  this  applies  only  to  the  extent  that 
interspecific  hybridization  is  excluded  or  non- 
introgressive.  Success  in  application  of  this 
species  definition  to  the  recognition  of  contem- 
porary species  and,  therefore,  the  "objective" 
bases  of  taxa  recognized  depend  on  the  degree 
to  which  reproductive  isolation  can  be  demon- 
strated or  inferred  where  it  exists  and  excluded 
where  it  does  not.  Direct,  unequivocal  evidence 
for  this  intrinsic  isolation  or  lack  of  same  can 
seldom,  if  ever,  be  provided,  even  from  breeding 
tests.  Only  criteria  by  which  indirect  evidence 
can  be  evaluated  are  generally  useful. 

The  following  working  criteria  were  used  in 
recognizing  species.  Two  sympatric  or  parapa- 
tric  forms  were  considered  separate  species  if 
all  individuals  (of  one  or  both  sexes)  of  one  differ 
from  all  members  (of  the  same  or  both  sexes)  of 
the  other  in  one  or  more  structural  characters 
(other  than  color  or  size).  In  practice,  members 
of  sympatric  or  parapatric  Nebria  species  gen- 
erally differ  in  several  characters  and  therefore 
easily  fulfill  this  criterion.  Two  allopatric  forms 
were  considered  separate  species  if  differences 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


between  members  of  each  are  roughly  equiva- 
lent to  differences  between  members  of  two 
closely  related  sympatric  species  AND  if  mem- 
bers of  geographically  intermediate  populations 
of  either  fail  to  exhibit  intermediate  character 
states  for  the  differentiating  characters.  Use  of 
the  above  criteria  required  the  following  as- 
sumption: that  continuity  or  discontinuity  in 
characters  of  external  structure,  internal  geni- 
talic  structure,  geographical  and  habitat  distri- 
bution, and  life  history  provided  by  the  study  of 
population  samples  (comprised  of  individuals 
and  groups  of  same)  is  evidence  of  reproductive 
continuity  or  isolation,  respectively,  among  nat- 
ural populations.  Clearly,  recognition  of  allo- 
patric  species  is  more  arbitrary  than  distinguish- 
ing sympatric  or  parapatric  species;  but 
hypotheses  about  relationship  must  be  drawn 
from  available  evidence,  and  some  estimate  of 
these  relationships  is  preferable  to  none. 

I  accept  the  subspecies  definition  of  Edwards 
(1956b:230)  in  slightly  altered  form;  namely,  that 
subspecies  are  populations  or  groups  of  popu- 
lations whose  members  are  recognizably  differ- 
ent from  members  of  other  conspecific  popula- 
tions and  would  interbreed  with  the  latter  IF  they 
occurred  sympatrically  and  synchronously  (un- 
der natural  conditions)  BUT  are  distinctly  iso- 
lated from  them  during  their  mating  periods. 

THE  SUBSPECIES  CATEGORY. — I  frequently 
used  the  subspecies  category  in  ranking  allopat- 
ric  Nebria  populations  and  population  groups 
because  patterns  of  geographical  variation  are 
often  such  that  discontinuities  in  structural  and 
other  characters  correlate  well  with  discontinu- 
ities in  habitat  or  physiography  (see  also  Ball 
1966;  and  Ball  and  Negre  1972).  Other  workers 
(Erwin  1970;  Madge  1967;  Noonan  1973;  and 
Whitehead  1976)  have  avoided  use  of  the  sub- 
species category  in  their  studies  of  lowland, 
mainland  groups.  Recognition  of  the  usefulness 
of  a  subspecies  concept  appears  to  depend  on 
one's  particular  perspective;  more  specifically, 
on  the  distribution  of  habitats  or  areas  occupied 
by  the  organisms  one  studies.  Where  gaps  be- 
tween areas  of  suitable  habitat  are  broad  and 
clear,  such  as  with  predominantly  montane 
groups  like  Nebria,  correlated  discontinuity  in 
variation  is  more  confidently  recognized  and 
suggestive  of  active,  effective  barriers  to  gene 
flow. 

Because  names  for  subspecies  are  accorded 
the  same  rights  and  strictures  as  species  names 


and  compete  with  the  latter  for  priority  under 
the  present  International  Code  of  Zoological 
Nomenclature,  the  description  of  new  subspe- 
cies (and  resultant  introduction  of  new  names 
into  the  literature)  must  be  justified.  I  suggest 
that  recognition  of  subspecies  as  defined  above 
is  justified  because  they  represent  incipient 
species.  They  are  at  present  independent  evo- 
lutionary units  whose  members  are  already  dif- 
ferentiated from  members  of  other  conspecific 
populations.  Because  these  units  are  generally 
of  zoogeographic  and  historical  significance,  the 
formal  subspecies  trinomen  is  as  welcome  a 
"shorthand  notation"  (Larson  1975:252)  as  the 
species  binomen  or  any  other  formal  name. 

In  addition  to  the  question  of  usefulness  is  that 
of  practicality,  recognizing  inherent  difficulties 
in  the  formulation  and  application  of  a  subspe- 
cies concept.  Use  of  the  subspecies  category  has 
been  warmly  debated  for  decades  (see  for  ex- 
ample, Brown  and  Wilson  1954;  Edwards  1954, 
1956a,  1956b;  Inger  1961;  and  Wilson  and  Brown 
1953).  Without  question,  great  variation  in  sub- 
species concepts  has  existed  among  taxono- 
mists,  so  that  taxa  recognized  by  various  work- 
ers are  seldom  equivalent  or  even  comparable 
units.  The  inability  of  taxonomists  to  formulate 
and  apply  a  common  subspecies  concept  to  their 
studies  is  unfortunate  but  understandable  in  light 
of  evolutionary  theory,  which  predicts  a  spec- 
trum of  intermediate  stages  in  the  process  of 
speciation.  Assignment  of  formal  subspecific 
rank  to  populations  at  one  particular  stage  in  the 
process  is  complicated  by  the  potential  for  al- 
most infinite  variety  in  evolutionary  rate  and 
direction  of  change. 

I  agree  with  Edwards  (1954,  1956a,  1956b)  that 
the  subspecies  as  defined  above  is  a  relatively 
objective  ("comparatively  but  not  absolutely 
nonarbitrary"  [Simpson  1961:116])  category. 
Extensive  barriers  to  gene  flow  between  sub- 
species can,  as  suggested  earlier,  be  recognized 
in  some  instances  through  detailed  studies  of 
habitat  and  geographical  distributions  and  life 
histories.  These  may  in  fact  be  easier  to  dem- 
onstrate than  intrinsic  isolating  mechanisms 
(i.e.,  reproductive  isolation)  between  separate 
species.  Arbitrariness  arises  in  decisions  on 
"how  different"  members  of  allopatric  or  al- 
lochronous  populations  must  be  for  recognition 
as  either  distinct  species  or  conspecific  subspe- 
cies. 

Working  criteria  I  used  for  recognizing  sub- 


94 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


species  were  as  follows.  Two  populations  or 
groups  of  populations  were  considered  separate 
subspecies  if  (1)  their  respective  geographical 
ranges  are  allopatric  and  separated  by  recogniz- 
able environmental  barriers,  (2)  differences  be- 
tween members  of  each  are  constant  but  less 
distinctive  than  differences  between  members  of 
two  closely  related  sympatric  species  in  one  or 
more  characters  of  structure,  color,  or  size,  AND 
(3)  the  geographical  pattern  of  variation  in  dis- 
tinguishing characters  is  nonclinal  or  distinctly 
step-clinal.  In  general,  all  members  of  a  subspe- 
cies can  be  recognized  on  structural  characters 
alone.  However,  in  some  instances,  where  two 
subspecies  were  recognized  as  distinct  by  a  step- 
clinal  pattern  of  variation,  all  members  of  one 
subspecies  may  be  distinguished  only  from  all 
members  of  the  most  proximate  populations  of 
the  other.  A  good  example  of  this  type  of  vari- 
ation pattern  is  seen  for  Nebria  lacustris  Casey 
(see  below;  details  to  be  presented  elsewhere 
[Kavanaugh,  manuscript  in  preparation]).  The 
following  assumption  was  required  for  use  of  the 
above  criteria:  that  the  amount  of  phenotypic 
divergence  between  members  of  two  allopatric 
populations  is  a  measure  not  only  of  the  com- 
pleteness and  duration  of  their  extrinsic  isolation 
but  also  of  the  probability  that  reproductive  (in- 
trinsic) isolation  has  evolved  between  them. 
Where  this  assumption  is  invalid,  the  resultant 
error  in  ranking  (i.e.,  recognizing  as  subspecies 
taxa  which,  in  fact,  conform  to  the  biological 
species  definition)  is,  in  my  opinion,  less  objec- 
tionable than  its  opposite.  If  two  de  facto 
species  are  ranked  as  subspecies,  close  relation- 
ship between  the  taxa  is  merely  overempha- 
sized, and  species-group  names  are  still  provid- 
ed and  protected  by  priority.  Only  changes  in 
status  are  required  if  actual  relationships  are 
correctly  determined  later. 

NEW  NEARCTIC  NEBRIA  SPECIES  AND 
SUBSPECIES 

The  order  of  presentation  of  new  taxa  in  this 
section  follows  a  new  classification  of  Nearctic 
Nebria  to  be  presented  elsewhere  (Kavanaugh, 
manuscript  in  preparation). 

Nebria  gouleti,  new  species 

(Figures  2,  13,  51) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wash.,  Asotin 
Co.,  Hwy.  129,  10  mi.  sw.  Antone,  Rattlesnake  Cr.,  3000',  25 
August  1973  DHKavanaugh  Family"/  "D.  H.  Kavanaugh  Col- 


lection" [orange  label]/  "Holotype  Nebria  gouleti  Kavanaugh 
det.  D.  H.  Kavanaugh  1976"  [red  label]/  "California  Academy 
of  Sciences  Type  No.  12504."  ALLOT YPE  (same  data  as  ho- 
lotype)  also  in  CAS.  In  total,  514  PARATYPES  (227  males  and 
288  females)  are  deposited  in  the  following  collections: 
AMNH,  ANSP,  BFCa,  CAS,  CUIC,  DHKa,  DJLa,  LRus, 
MCZ,  MSU,  OSUO,  PURC,  RTBe,  SJSC,  UASM,  UIMI, 
UMMZ,  USNM,  UWBM,  and  WSU. 

TYPE- LOCALITY. — Rattlesnake  Creek  (10  miles  [ca.  16  km] 
sw  of  Antone),  Asotin  County,  Washington. 

DIAGNOSTIC  COMBINATION. — Head  uniform- 
ly dark,  without  pale  spot(s)  on  vertex,  antennal 
scape  moderately  long,  symmetrically  ellipsoid 
(Fig.  2);  pronotum  distinctly  cordate,  midlateral 
seta  present  (Fig.  13),  without  longitudinal  tu- 
bercle medial  to  midlateral  seta;  elytra  dull,  mi- 
crosculpture  deeply  impressed,  meshes  isodia- 
metric  or  very  slightly  transverse,  fifth  elytral 
interval  without  setae;  metepisternum  impunc- 
tate;  hind  coxa  bi-  or  trisetose  basally,  tarso- 
meres  of  hind  tarsus  glabrous  dorsally;  second 
visible  abdominal  sternum  with  patch  of  setae 
medially  (between  hind  coxae),  third  to  fifth  vis- 
ible sterna  each  with  two  to  six  pairs  of  posterior 
paramedial  setae  (Fig.  37). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  species  in  honor  of  my 
good  friend  and  frequent  field  companion,  Henri 
Goulet,  from  whom  I  have  learned  to  observe 
and  appreciate  carabid  beetles  while  they  are 
still  alive! 

GEOGRAPHICAL  DISTRIBUTION. — Figure  51; 
restricted  to  portions  of  the  Columbia  Plateau 
region  in  Washington,  Oregon,  and  Idaho;  the 
lower  Columbia  River  valley;  and  the  mountains 
of  northern  and  central  Idaho.  I  have  studied 
specimens  from  the  following  localities. 

United  States  of  America 

IDAHO:  Adams  County,  New  Meadows  ([1,170  m])  [June] 
(1;  UIMI);  Bonner  County,  Sandpoint  [Sep.]  (1;  CAS);  Clear- 
water  County,  Canyon  Ranger  Station  [Sep.]  (1;  UIMI);  Cus- 
ter  County,  Lower  Stanley  (10  miles  [ca.  16  km]  E  on  Salmon 
River  [1,800m])  [Aug.]  (1:  DHKa),  Salmon  River  (at  Bayhorse 
Creek  [1,620  m])  [Aug.]  (45;  DHKa);  Idaho  County,  Fenn 
Ranger  Station  (Selway  Fork  Clearwater  River)  [Aug.]  (3; 
UIMI),  Lochsa  River  (at  Canyon  Creek  [520  m])  [July]  (2; 
DHKa),  Lowell  (39  miles  [ca.  63  km]  NE  on  Lochsa  River  [850 
m])  [July]  (1;  DHKa),  Meadow  Creek  (1.5  miles  [ca.  2.4  km] 
s  of  Selway  Falls)  [Sep.]  (I;  CAS),  Orogrande  Creek  [Aug.] 
(3;  UWBM),  Selway  Falls  [July]  (1:  UWBM):  Kootenai  Coun- 
ty, Coeur  d'Alene  [June-July]  (5;  CAS,  MCZ,  USNM),  Hay- 
den  Lake  [Aug.]  (9;  AMNH,  MCZ,  UIMI,  USNM):  Latah 
County,  Kendrick  [Sep.]  (2:  UIMI),  Moscow  (1;  USNM); 
Lemhi  County,  Salmon  (and  21  miles  [ca.  34  km]  N)  [July- 
Aug.]  (2;  UASM,  UIMI);  Nez  Perce  County,  Lenore  [May] 
(2;  UIMI),  Lewiston  (and  2  and  3  miles  [ca.  3.2  and  4.8  km] 


KAVANAUGH:  NEW  NEARCTIC  NEBR1A 


E  on  Clearwater  River  [300  m-380  m])  [May-Aug.,  Oct.]  (52; 
DHKa.  UIMI),  Waha  [Aug.]  (5;  CAS,  MCZ,  PURC,  USNM), 
Waha  Lake  [July]  (4;  CAS,  MCZ);  Shoshone  County,  Avery 
[Sep.]  (2;  UWBM),  Clarkia  [June]  (1;  UIMI),  St.  Joe  River 
(17  miles  [ca.  27  km]  E  of  St.  Maries)  [July]  (10;  CAS);  Valley 
County,  Dagger  Falls  [July]  (2:  BFCa),  Zena  Creek  [June]  (1; 
UIMI);  County  unknown.  Bungalow  [Aug.]  (15;  UWBM). 
OREGON:  Baker  County,  East  Fork  Eagle  Creek  [July]  (2; 
UWBM),  Halfway  [July]  (1;  BFCa);  Hood  River  County, 
Hood  River  [Sep.]  (2;  MCZ);  Multnomah  County,  Portland 
[July]  (3;  CUIC);  Umatilla  County,  Cottonwood  Creek  [June] 
(3;  USNM,  UWBM),  Kamela  [June]  (1;  CAS),  Milton  (Walla 
Walla  River)  [May-June]  (4;  UWBM),  s  of  Kooskooskie 
(Washington)  [June]  (2;  UWBM);  Union  County,  Elgin  (and 
0.1  mile  [0.16  km]  s  at  Grande  Ronde  River  [1,130  m])  [May, 
Aug.]  (2;  DHKa),  Hilgard  Junction  State  Park  [Aug.]  (3; 
RTBe),  La  Grande  (1  mile  [ca.  1.6  km]  NE  on  Grande  Ronde 
River  [1,070  m])  [May]  (5;  DHKa);  Wallowa  County,  Minam 
(0.2  miles  [ca.  0.3  km]  SE  at  junction  of  Minam  and  Wallowa 
rivers  [790  m])  [May]  (5;  DHKa).  WASHINGTON:  Asotin  Coun- 
ty, Antone  ( 10  miles  [ca.  16  km]  sw  at  Rattlesnake  Creek  [910 
m])  [Aug.]  (21;  DHKa),  Asotin  (Asotin  Creek  [370  m])  [May- 
June]  (18;  ANSP,  CAS,  DHKa),  Buford  Creek  (3.6  miles  [ca. 
5.8  km]  s  of  Grande  Ronde  River  [760  m])  [May]  (8;  DHKa), 
Grande  Ronde  River  (at  Highway  129  [550  m])  [May,  Sep.] 
(2;  DHKa,  SJSC);  Chelan  County,  Leavenworth  [July]  (20; 
ANSP,  CAS,  MCZ,  PURC,  USNM,  UWBM),  Peshastin 
Creek  [July]  (1;  UASM),  Tumwater  Canyon  ([610  m])  [May] 
(2;  LRus);  Columbia  County,  Dayton  (Touchet  River  [580  m]) 
[May,  July]  (5:  DKHa,  WSU);  Franklin  County,  Palouse  Falls 
State  Park  [May]  (2;  LRus);  Klickitat  County,  Trout  Lake 
[July]  (1;  UWBM),  White  Salmon  [Aug.]  (2;  UWBM);  Ska- 
mania  County,  Underwood  (15  miles  [ca.  24  km]  w  on  Colum- 
bia River  [90  m])  [May]  (4;  DHKa);  Spokane  County,  Spokane 
Falls  (1;  CAS):  Walla  Walla  County,  Burbank  [May]  (1; 
ANSP),  Coppei  [May]  (1:  UAFA),  Coppei  Creek  [May]  (1; 
UIMI),  Kooskooskie  [June,  Aug.]  (33;  UIMI,  UWBM), 
Touchet  River  (near  Waitsburg)  [July]  (98;  DJLa),  Walla  Wal- 
la (and  2  miles  [ca.  3.2  km]  s  [320  m],  Mill  Creek)  [May-Sep.] 
(139;  CAS.  CUIC,  DHKa,  MCZ,  OSUO,  PURC,  UIMI, 
UMMZ,  UWBM,  WSU);  Whitman  County,  Almota  [May]  (2; 
MCZ,  MSU),  Wawawai  (2.8  miles  [ca.  4.5  km]  NW  at  Snake 
River  [300  m])  [May]  (11;  DHKa),  Wilma  [Apr.-May]  (6; 
ANSP,  UWBM). 

Doubtful  Records 

CANADA — ALBERTA:  Longview  (Highwood  River)  [July] 
(7:  CAS).  UNITED  STATES  OF  AMERICA- — CALIFORNIA:  (2; 

USNM). 

Nebria  lacustris  bellorum,  new  subspecies 

(Figures  14,  52) 

Nebria  lacustris:  BELL  1955:265  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Tenn..  Great 
Smoky  Mts.  N.  P.,  Chimneys  Picnic  Area,  W.  Prong  Little 
Pigeon  R.,  3000',  24May73  DHKavanaugh  &  HGoulet'V  "D. 
H.  Kavanaugh  Collection"  [orange  label]/  "Holotype  Nebria 
lacustris  bellorum  Kavanaugh  det.  D.  H.  Kavanaugh  1976" 
[red  label]/  "California  Academy  of  Sciences  Type  No. 
12506."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In 
total,  276  PARATYPES  (139  males  and  137  females)  are  depos- 
ited in  the  following  collections:  CAS,  CUIC,  DHKa,  HoKn. 
MCZ,  RTBe,  TCBa,  UAFA,  UASM,  USNM. 


TYPE-LOCALITY. — West  Prong  Little  Pigeon  River  (at 
Chimneys  Picnic  Area),  Great  Smoky  Mountains  National 
Park,  Tennessee. 

DIAGNOSTIC  COMBINATION. — Head  uniform- 
ly dark,  without  pale  spot(s)  on  vertex;  prono- 
tum  (Fig.  14)  distinctly  cordate,  midlateral  seta 
present,  without  longitudinal  tubercle  medial  to 
midlateral  seta;  elytra  very  shiny  or  faintly  iri- 
descent, microsculpture  consists  of  thin,  trans- 
verse lines  without  (or  with  only  scattered) 
meshes,  striae  very  deeply  impressed,  intervals 
markedly  convex,  fifth  interval  without  setae; 
metepisternum  impunctate;  tarsomeres  of  hind 
tarsus  glabrous  dorsally;  second  visible  abdom- 
inal sternum  glabrous  medially,  third  to  fifth  vis- 
ible sterna  each  with  single  pair  of  posterior  par- 
amedial  setae;  specimen  from  locality  in 
southeastern  United  States,  south  of  French 
Broad  River  (North  Carolina  or  Tennessee)  (Fig. 
52). 

DERIVATION  OF  TAXON  NAME. — I  take  plea- 
sure in  naming  this  subspecies  in  honor  of  Ross 
and  Joyce  Bell  (Burlington,  Vermont),  who  have 
long  been  interested  in  the  Nebria  of  eastern 
North  America,  and  who  have  generously  pro- 
vided encouragement  and  information  for  this 
study. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  52; 
restricted  to  that  portion  of  the  southern  Appa- 
lachian region  including  Great  Smoky  Moun- 
tains National  Park  and  adjacent  mountain 
ranges.  I  have  studied  specimens  from  the  fol- 
lowing localities. 

United  States  of  America 

NORTH  CAROLINA:  Great  Smoky  Mountains  National  Park 
[June]  (1;  MCZ),  Indian  Gap  [Sep.]  (1;  USNM),  Kephard 
Prong  Oconaluftee  River  [Aug.]  (14;  DHKa,  TCBa),  New- 
found Gap  ([1,520  m-1,580  m])  [Aug.]  (1:  MCZ),  Oconaluftee 
River  (1 1  miles  [ca.  18  km]  SE  of  Newfound  Gap  on  Highway 
441  [550  m],  at  Smokemont  [840  m]  and  2.3  miles  [ca.  3.7  km] 
NW  of  Smokemont  [610  m])  [May,  Sep.]  (61;  DHKa,  USNM), 
Smokemont  Loop  Trail  ([760  m-910  m])  [July]  (18;  UAFA); 
Haywood  County,  Beach  Gap  (7  miles  [ca.  11  km]  s  of  Sun- 
burst at  Middle  Prong  West  Fork  Pigeon  River  [1,520  m]) 
[June,  Aug.]  (4;  HoKn),  Bubbling  Spring  Branch  ([1,520  m]) 
[July]  (10;  DHKa,  TCBa),  Graveyard  Fields  (Blue  Ridge  Park- 
way at  Yellowstone  Prong  Pigeon  River  [1,540  m])  [May]  (1: 
DHKa),  Mount  Sterling  ([910  m])  [Oct.]  (7;  CAS,  CUIC),  Re- 
treat [May]  (6:  USNM);  Swain  County,  Bryson  City  (Deep 
Creek  [610  m])  [Aug.]  (2;  MCZ).  TENNESSEE:  Great  Smoky 
Mountains  National  Park,  Alum  Cave  Creek  ([1,220  m-1,250 
m])  [May]  (4;  DHKa),  Chimneys  Camp  ([910  m])  [May,  Sep.] 
(18;  CAS,  CUIC,  USNM),  Newfound  Gap  ([1,520  m-1,580 
m])  [Aug.]  (2:  MCZ,  UASM),  Tennessee  Branch  Bradley 
Creek  ([980  m])  [May]  (15;  USNM),  Walker  Prong  Little  Pi- 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


geon  River  [Aug.]  (1;  DHKa),  West  Prong  Little  Pigeon  River 
([1,040  m-1,220  m]),  Chimneys  Picnic  Area  [910  m],  (3  miles 
[ca.  4.8  km]  s  of  Gatlinburg  [520  m],  3.7  miles  [ca.  6.0  km]  N 
of  Newfound  Gap  [1,220  m])  [May,  July,  Sep.]  (109;  DHKa, 
MCZ,  UAFA,  USNM);  Sevier  County,  Gatlinburg  [June]  (1; 
DHKa). 

Doubtful  Records 

UNITED  STATES  OF  AMERICA — COLORADO:  Jefferson 
County,  Golden  (Clear  Creek)  [June]  (1;  USNM).  LOUISIANA: 
St.  Tammany  Parish,  Highway  90  (10  miles  [ca.  16  km]  w  of 
Mississippi  border)  [June]  (1;  CAS). 

Nebria  nivalis  gaspesiana,  new  subspecies 

(Figures  15,  53) 

Nebria  suturalis;  LENG  (not  LeConte).  — LENG  1920:47  (in 

part).  — LINDROTH  1955a:39. 
Nebria  nivalis  bifaria  MANNERHEIM.  — LINDROTH  1955a:39 

(in  part). 
Nebria  nivalis;  auctorum.  — LAROCHELLE  1972:62  (in  part); 

1976:48;  1977:12  (in  part).  —LINDROTH  1961:81  (in  part); 

1971:1459. 
Nehria  bifaria;  LINDROTH  1961:81  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "Can.,  Que.,  Penin- 
sule  de  Gaspe,  Mt.  Albert,  Ruisseau  du  Diable,  3200'-3500', 
11-12  June  73  DHKavanaugh  &  HGoulet'V  "D.  H.  Kavan- 
augh  Collection"  [orange  label]/  "Holotype  Nebria  nivalis 
gaspesiana  Kavanaugh  det.  D.  H.  Kavanaugh  1976"  [red  la- 
bel]/ "California  Academy  of  Sciences  Type  No.  12510."  AL- 
LOTYPE  (same  data  as  holotype)  also  in  CAS.  In  total,  287 
PARATYPES  (119  males  and  168  females)  are  deposited  in  the 
following  collections:  ALar,  CAS,  CNC,  DBUM,  DHKa, 
MCZ,  RTBe,  UASM,  USNM. 

TYPE-LOCALITY.— Ruisseau  du  Diable  (980  m-1,070  m), 
Mont  Albert,  Pare  de  La  Gaspesie,  Comte  de  Gaspe-Ouest, 
Quebec. 

DIAGNOSTIC  COMBINATION. — Head  uniform- 
ly dark,  without  pale  spot(s)  on  vertex;  prono- 
tum  (Fig.  15)  distinctly  cordate,  midlateral  seta 
present,  with  small  longitudinal  tubercle  medial 
to  midlateral  seta;  fifth  elytral  interval  with  one 
to  four  setae;  metepisternum  impunctate;  tar- 
someres  of  hind  tarsus  glabrous  dorsally;  spec- 
imen from  locality  in  Maine  (Mount  Katahdin), 
Gaspe  Peninsula,  Island  of  Newfoundland,  or 
Labrador  (west  only  to  east  shore  of  Ungava 
Bay)  (Fig.  53). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Gaspe  Peninsula,  on  which 
the  type-locality  is  found. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  53; 
disjunct;  restricted  to  the  western  part  of  the 
Island  of  Newfoundland,  Mount  Katahdin 
(Maine),  the  high  mountains  of  the  Gaspe  Pen- 
insula, and  coastal  Labrador  (northwest  to  the 
eastern  shore  of  Ungava  Bay).  I  have  studied 
specimens  from  the  following  localities. 


Canada 

NEWFOUNDLAND:  Island  of  Newfoundland,  Little  Codroy 
River  [July]  (4;  MCZ,  USNM),  St.  John  Bay  (Doctors  Brook) 
[July]  (5;  CNC,  ZMLS),  South  Branch  (Grand  Codroy  River) 
[July]  (4;  MCZ),  Spruce  Brook  (1;  MCZ).  QUEBEC:  Comte  de 
Bonaventure  [July]  (1;  ALar);  Comte  de  Gaspe-Ouest,  Pare 
de  La  Gaspesie  (Mont  Albert  at  Ruisseau  du  Diable  [980  m- 
1,070  m])  [June-July]  (261;  ALar,  CNC,  DBUM,  DHKa, 
RTBe);  Territoire  du  Nouveau-Quebec,  Riviere  Abloviak 
[July](l;  DBUM). 

United  States  of  America 

MAINE:  Piscataquis  County,  Baxter  State  Park  (Mount  Ka- 
tahdin [910  m-1,580  m])  [June-Aug.]  (10;  CAS,  MCZ, 

USNM). 

Nebria  gyllenhali  lassenensis,  new  subspecies 

(Figures  3,  54) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Calif.,  Las- 
sen  Volcanic  N.  P.,  Cascade  Range,  s.  slope  Mt.  Lassen, 
Emerald  Lk.,  2450m,  22Aug.74  D.H. Kavanaugh  Coil."/  "D. 
H.  Kavanaugh  Collection"  [orange  label]/  "Holotype  Nebria 
gyllenhali  lassenensis  Kavanaugh  det.  D.  H.  Kavanaugh 
1976"  [red  label]/  "California  Academy  of  Sciences  Type  No. 
12511."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In 
total,  34  PARATYPES  (18  males  and  16  females)  are  deposited 
in  the  following  collections:  CAS,  DHKa,  FMNH,  USNM. 

TYPE-LOCALITY. — Emerald  Lake,  Mount  Lassen  (south 
slope),  Lassen  Volcanic  National  Park,  California. 

DIAGNOSTIC  COMBINATION. — Head,  prono- 
tum,  and  elytra  without  metallic  reflection;  head 
uniformly  dark,  without  pale  spot(s)  on  vertex, 
moderate  in  size  and  convexity,  with  genae  and 
occiput  not  broadened,  eyes  full-sized;  antennal 
scape  (Fig.  3)  short,  straight,  symmetrically 
ovoid  or  anteriorly  more  convex;  penultimate 
labial  palpomere  trisetose;  pronotum  small, 
slightly  narrow,  moderately  cordate  (lateral 
margin  moderately  arcuate  at  middle),  slight- 
ly dull  to  moderately  shiny,  microsculpture 
moderately  to  faintly  impressed,  meshes  isodi- 
ametric,  alveolae  flat,  midlateral  seta  present, 
without  longitudinal  tubercle  medial  to  midlat- 
eral seta;  elytral  intervals  flat,  third  interval  con- 
tinuous or  faintly  catenate  (setiferous  pores 
slightly  to  moderately  foveate),  fifth  interval 
without  setae;  hindwing  full-sized;  metepister- 
num impunctate;  tarsomeres  of  hind  tarsus  gla- 
brous dorsally;  second  visible  abdominal  ster- 
num glabrous  medially,  third  to  fifth  visible 
sterna  each  with  single  pair  of  posterior  para- 
medial  setae;  specimen  from  locality  in  northern 
Sierra  Nevada  (California)  or  Cascade  Range 
south  of  Columbia  River  (Fig.  54). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


97 


cies  is  named  for  Mount  Lassen,  on  the  south 
slope  of  which  the  type-locality  is  found. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  54; 
restricted  to  the  southern  Cascade  Range  (south 
of  the  lower  Columbia  River  valley)  and  the 
northern  Sierra  Nevada  (south  to  Sonora  Pass). 
I  have  studied  specimens  from  the  following  lo- 
calities. 

United  States  of  America 

CALIFORNIA:  Lassen  Volcanic  National  Park,  Emerald 
Lake  ([2,450  m])  [Aug.,  Oct.]  (6;  CAS,  DHKa,  FMNH),  Little 
Hot  Springs  Area  (East  Sulphur  Creek  [2,440  m-2,530  m]) 
[Aug.]  (6;  CAS,  DHKa),  Mount  Lassen  (3  miles  [ca.  4.8  km] 
SE)  [July]  (1;  USNM),  Ridge  Lake  [Aug.]  (16;  CAS);  Tuol- 
umne  County,  Sonora  Pass  [Aug.]  (3;  CAS).  OREGON:  Des- 
chutes  County,  Todd  Lake  (and  1  mile  [ca.  1.6  km]  s)  [Aug.] 
(4;  JSch). 

Nebria  gyllenhali  lindrothi,  new  subspecies 

(Figures  4,  54) 

Nebria  sahlbergi;  WICKHAM  1902:232. 

Nebria  gyllenhali;  auctorum.  — ARMIN  1963:90. 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wyo.,  Albany 
Co.,  Medicine  Bow  Mts.,  Snowy  Range,  Brooklyn  Lk,  10500', 
HJuly72  DHKavanaugh'V  "D.  H.  Kavanaugh  Collection" 
[orange  label]/  "Holotype  Nebria  gyllenhali  lindrothi  Kavan- 
augh det.  D.  H.  Kavanaugh  1976"  [red  label]/  "California 
Academy  of  Sciences  Type  No.  12512."  ALLOTYPE  (same 
data  as  holotype)  also  in  CAS.  In  total,  1579  PARATYPES  (795 
males  and  784  females)  are  deposited  in  the  following  collec- 
tions: AMNH,  ANSP,  CArm,  CAS,  CDA,  CNC,  CUB, 
CUIC,  DHKa,  DRWh,  EAMa,  FMNH,  ICCM,  KSUC,  MCZ, 
MSU,  PADA,  PMCh,  PMNH,  PURC,  RCGr,  RDav,  RTBe, 
SDSU,  UAFA,  UASM,  USNM,  UWBM,  UWL,  ZMLS. 

TYPE-LOCALITY. — Brooklyn  Lake  (3,200  m),  Albany  Coun- 
ty, Wyoming. 

NOTES  ON  NOMENCLATURE  AND  TYPES. — 
This  subspecies  is  the  form  noted  by  Lindroth 
(1969b:1109)  as  an  undescribed  species  related 
to  N.  gyllenhali. 

DIAGNOSTIC  COMBINATION. — Head,  prono- 
tum,  and  elytra  without  metallic  reflection;  head 
uniformly  dark,  without  pale  spot(s)  on  vertex, 
moderate  in  size  and  convexity,  with  genae  and 
occiput  not  broadened,  eyes  full-sized;  antennal 
scape  (Fig.  4)  long,  straight,  slightly  arcuate, 
moderately  or  markedly  narrowed  basally;  pen- 
ultimate labial  palpomere  trisetose;  pronotum 
small,  slightly  narrow,  moderately  cordate  (lat- 
eral margin  moderately  arcuate  at  middle), 
slightly  dull  to  moderately  shiny,  microsculpture 
moderately  to  faintly  impressed,  meshes  isodi- 
ametric,  alveolae  flat,  midlateral  seta  present, 
without  longitudinal  tubercle  medial  to  midlat- 


eral seta;  elytral  intervals  flat,  third  interval  con- 
tinuous or  faintly  catenate  (setiferous  pores 
slightly  to  moderately  foveate),  fifth  interval 
without  setae;  hind  wing  reduced  in  size  and  ve- 
nation pattern;  metepisternum  impunctate;  tar- 
someres  of  hind  tarsus  glabrous  dorsally;  second 
visible  abdominal  sternum  glabrous  medially, 
third  to  fifth  visible  sterna  each  with  single  pair 
of  posterior  paramedial  setae;  specimen  from  lo- 
cality in  southern  Rocky  Mountains — in  Medi- 
cine Bow  Mountains  or  Sierra  Madre  (Wyo- 
ming), or  Uinta  Mountains  (Utah),  or  further 
south  (Fig.  54). 

DERIVATION  OF  TAXON  NAME. — I  am  hon- 
ored to  name  this  subspecies  after  Carl  H.  Lind- 
roth, a  friend  whose  knowledge  of  and  studies 
on  Holarctic  Carabidae,  and  Nebria  in  particu- 
lar, have  been  a  guiding  light  in  this  study,  and 
who  first  called  my  attention  to  the  distinctive- 
ness  of  members  of  this  taxon.  I  deeply  regret 
that  Carl  did  not  live  to  see  the  completion  of 
this  study,  to  which  he  contributed  so  much. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  54; 
restricted  to  the  southern  Rocky  Mountains, 
from  northern  New  Mexico  north  to  the  Medi- 
cine Bow  Mountains  and  Sierra  Madre  of  south- 
ern Wyoming,  west  to  the  Uinta  Mountains  of 
eastern  Utah.  I  have  studied  specimens  from  the 
following  localities. 

United  States  of  America 

COLORADO:  (2;  KUSM,  PURC);  Boulder  County,  ([3,810 
m])  [July]  (1;  CUB),  Arapaho  Pass  ([2,740  m-3,350  m])  [Aug.] 
(1;  RTBe),  Arapaho  Peak  ([3,510  m])  [Aug.]  (3;  CArm),  Bea- 
ver Creek  ([2,530  m])  [July]  (1;  CArm),  Fourth  of  July  Mines 
([3,430  m])  [Aug.]  (2;  CArm),  Jenny  Lake  ([3,200  m])  [July] 
(2;  CArm),  Middle  St.  Vrain  Creek  ([3,600  m])  [July]  (1; 
CArm),  Niwot  Ridge  ([3,050  m])  [July]  (5;  CArm,  CUB),  Rain- 
bow Lakes  ([3,350  m])  [Aug.]  (3;  CNC),  Rollins  Pass  ([3,410 
m-3,540  m])  [Aug.]  (2;  CUB,  DHKa),  South  Fork  Middle 
Boulder  Creek  ([2,900  m-3,960  m])  [Aug.]  (1;  CArm);  Clear 
Creek  County,  Leavenworth  Valley  (Argentine  Road  [3,660 
m-3,960  m])  (2;  USNM),  Loveland  Pass  ([3,660  m-3,840  m]) 
[July]  (42;  CAS,  RTBe,  UWBM),  Mount  Evans  ([3,600  m- 
4,330  m],  Summit  Lake  [3,900  m-3,960  m])  [July-Aug.J  (76; 
CAS,  CNC,  DHKa,  FMNH,  MCZ,  PURC,  RCGr,  UASM); 
Conejos  County,  Cumbres  Pass  ([3,050  m])  [June]  (1;  CAS); 
Custer  County,  Ophir  Creek  (7.3  miles  [ca.  11.7  km]  sw  of 
Fairview  [3,140  m])  [June]  (9;  DHKa),  St.  Charles  River  (1.9 
miles  [ca.  3. 1  km]  sw  of  San  Isabel  [2,710  m-2,830  m])  [June] 
(14;  DHKa),  Wet  Mountains  ([3,470  m])  [July]  (2;  USNM); 
Dolores  County,  Lizard  Head  Pass  ([3,050  m])  [Sep.]  (1; 
CAS),  Rico  area  ([2,590  m-3,050  m])  [July]  (10:  ANSP,  MCZ, 
USNM);  El  Paso  County,  Pikes  Peak  ([3,960  m])  [July]  (8; 
AMNH,  CAS),  South  Cheyenne  Creek  (at  Gold  Camp  Road 
[2,530  m-2,650  m])  [June]  (1;  DHKa):  Garfield  County,  Trap- 
pers Lake  ([2,530  m-2,650  m])  [June]  (49;  DHKa);  Gilpin 


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County,  Central  City  ([2,620  m])  [July]  (1;  CAS),  Corona 
([3,350  m])  [Sep.]  (7;  CAS,  CUIC),  South  Boulder  Creek  (at 
Rollinsville  and  2.5  miles  [ca.  4.0  km]  w  [2,440  m])  [July- 
Aug.]  (5:  CArm,  DHKa,  UAFA);  Grand  County,  Ptarmigan 
Peak  (20:  MCZ,  ZMLS);  Gunnison  County,  Gothic  ([2,930  m]) 
[July]  (2:  USNM),  Mexican  Cut  Lakes  (1  mile  [ca.  1.6  km] 
NW  of  Elko  Park  [3,440  m])  [Aug.]  (1;  PMNH):  Huerfano 
County,  Apishapa  Pass  [June]  (3;  CUB),  Bear  Lake  [June] 
(13;  DHKa,  EAMa),  Blue  Lake  [June]  (10;  DHKa,  EAMa), 
Cucharas  Pass  ([3,050  m]  and  1.5  miles  [ca.  2.4  km]  N)  [June] 
(109;  DHKa,  EAMa),  Cucharas  River  (near  Blue  Lake  [3,190 
m])  [June]  (5;  DHKa);  Jackson  County,  Cameron  Pass  ([3,140 
m])  [Aug.]  (1;  UASM),  Rabbit  Ears  Pass  ([2,740  m])  [June- 
Aug.]  (8;  CAS,  USNM);  La  Plata  County,  Hermosa  Creek 
Trail  (NW  of  Hermosa  [2,960  m])  [Aug.]  (9;  UASM);  Larimer 
County,  Browns  Lake  Trail  [Aug.]  (9;  RTBe),  Crown  Point 
[July]  (21;  RTBe),  Zimmerman  Lake  [Aug.]  (3;  RTBe):  Los 
Animas  County,  Apishapa  Pass  (2.1  miles  [ca.  3.4  km]  SE 
[3,230  m])  [June]  (9;  DHKa),  Bear  Creek  (5  miles  [ca.  8  km] 
s  of  Cucharas  Pass  [2,800  m])  [June]  (83:  DHKa);  Mesa  Coun- 
ty [Aug.]  (5;  PMCh,  RDav),  Grand  Mesa  ([3,050  m])  [July- 
Aug.)  (6;  CAS,  CUB,  USNM),  Skyway  ([3,050  m])  [June]  (2; 
DHKa,  FMNH);  Mineral  County,  Pagosa  Springs  (20  miles 
[ca.  32  km]  N)  [June]  (1;  DRWh),  Wolf  Creek  Pass  ([3,250  m- 
3,660  m]  and  2  miles  [ca.  3.2  km]  w,  Thunder  Mountain  [3,570 
m])  [June-Aug.]  (96;  CAS,  CDA,  CNC,  DHKa,  EAMa, 
UASM,  USNM):  Ouray  County,  American  Flats  ([3,660  m]) 
[July]  (4;  AMNH),  Ouray  area  [June-July]  (4;  AMNH,  CAS); 
Park  County,  Middle  Fork  North  Platte  River  (near  Hoosier 
Pass  [3,540  m])  [July]  (1;  DHKa),  Mount  Lincoln  ([3,350  m]) 
[Aug.]  (1;  UASM);  Rio  Grande  County,  Rio  Grande  River  (1 
mile  [ca.  1.6  km]  s  of  South  Fork  [2,400  m])  [Aug.]  (29:  DHKa, 
EAMa),  Summitville  ([3,050  m-3,350  m])  [Aug.]  (7:  CNC, 
MCZ);  Rocky  Mountain  National  Park  ([3,200  m-3,510  m]) 
[June-Aug.]  (11;  CAS,  CNC,  UASM),  Fall  River  Pass  ([3,600 
m])  [July]  (5;  CUB),  Longs  Peak  ([3,840  m].  Chasm  Lake 
[3,810  m])  [July-Aug.]  (14;  CArm),  Milner  Pass  ([3,280  m]) 
[July]  (1;  AMNH),  Ouzel  Falls  ([3,050  m])  [Aug.]  (1;  CArm), 
Sandbeach  Lake  ([3,050  m])  [June]  (7:  CArm),  Trail  Ridge 
Road  ([3,660  m-3,960  m])  [Aug.]  (2;  CUB,  USNM);  Routt 
County,  Buffalo  Pass  (Summit  Lake  [3,140  m])  (4;  CAS),  Wal- 
ton Creek  (above  Dumont  Lake  [2,900  m-2,960  m])  [Aug.]  (5; 
DHKa);  San  Juan  County,  Silverton  ([3,660  m])  [July]  (1; 
USNM);  Summit  County,  Quandary  Peak  (south  slope  [3,410 
m-3,960  m])  [Aug.]  (35;  DHKa);  Teller  County  ([3,410  m]) 
[June]  (1:  PADA),  Phantom  Canyon  ([2,960  m])  [June]  (1; 
CAS);  County  unknown,  "Hot  Springs"  (1;  ICCM),  "King's 
Lake"  ([3,350  m])  [Aug.]  (2;  CArm),  La  Plata  Mountains 
([3,200  m-3,660  m])  [July-Aug.]  (4;  DHKa,  USNM),  "Lara- 
mie  Co."  (2;  ICCM).  Medicine  Bow  Range  ([2,440  m-3,660 
m])  [July]  (3;  MCZ),  Mount  Baldy  (summit  [3,750  m])  [July] 
(1;  CArm),  San  Juan  Mountains  ([3,660  m])  [July](l:  USNM), 
"foothills  of  Rocky  Mts."  (1;  PMNH).  NEW  MEXICO:  (9; 
ANSP,  ICCM,  KUSM,  MCZ,  USNM);  Mora  County,  Pecos 
Baldy  (2;  AMNH);  Rio  Arriba  County,  Gregorio  Lake  ([2,790 
m])  [June]  (5;  DHKa),  Rio  Puerco  (at  Rio  Puerco  Campground 
[2,350  m-2,440  m])  [June]  (3;  DHKa);  San  Miguel  County, 
Beaver  Creek  (sw  of  Rincon  Montoso  [2,620  m])  [Aug.]  (15; 
UASM),  Cowles  area  ([2,440  m],  Spirit  Lake  Trail)  [June]  (4; 
AMNH,  UASM),  Las  Vegas  area  ([2,130  m])  [July]  (1: 
KUSM),  Willow  Creek  (near  Torrero  [2,740  m])  [June]  (8; 
UASM);  Santa  Fe  County,  Lake  Katherine  ([3,600  m])  [June] 
(30;  DHKa),  Lake  Peak  (5;  AMNH),  Nambe  Divide  ([3,350 
m])  [July]  (2:  ANSP),  Santa  Fe  Ski  Basin  (5  miles  [ca.  8  km] 


NW)  (1;  AMNH);  Taos  County,  Red  River  (2.2  miles  [ca.  3.5 
km]  [2,610  m],  4.1  miles  [ca.  6.6  km]  [2,580  m],  and  5  miles 
[ca.  8  km]  w  of  Red  River,  8  miles  [ca.  13  km]  s  of  Red  River 
[2,870  m])  [June]  (40:  AMNH,  DHKa),  Tres  Ritos  [July]  (4; 
CAS),  Wheeler  Peak  ([3,050  m-3,640  m],  NW  slope.  Middle 
Fork  Lake  [3,340  m].  Middle  Fork  Red  River  [3,260  m-3,690 
m])  [June]  (172;  AMNH,  DHKa).  UTAH:  Duchesne  County, 
Garfield  Basin  (stream  below  Tungsten  Lake  [3,450  m])  [Sep.] 
(3;  USNM);  Grand  County,  Mill  Creek  (at  Oowah  Lake  [2,680 
m])  [Aug.]  (12;  DHKa);  San  Juan  County  ([2,800  m])  [Aug.] 
(1;  FMNH),  Dark  Canyon  ([2,960  m],  NE  slope  [3,240  m]) 
[Aug.]  (30:  UASM),  Geyser  Pass  ([3,050  m],  N  slope  [3,120 
m],  E  slope  [3,220  m],  at  Blue  Lake  [2,960  m])  [July-Aug.] 
(79;  AMNH,  UASM),  Mill  Creek  Valley  ([2,900  m])  [Aug.] 
(2;  UASM).  WYOMING:  Albany  County,  Brooklyn  Lake 
([3,200  m])  [July]  (41:  DHKa),  Centennial  area  ([3,050  m]) 
[Aug.]  (22:  MSU,  SDSU),  Douglas  Creek  (1  mile  [ca.  1.6  km] 
SSE  of  Keystone  [2,680  m])  [July]  (32;  DHKa),  Little  Brooklyn 
Lake  ([3,120  m])  [July]  (113:  DHKa),  Medicine  Bow  Peak 
([3,350  m-3,660  m])  [July-Aug.]  (3;  FMNH,  KSUC,  UWL), 
Mirror  Lake  ([3,290  m])  [July]  (37;  USNM),  Nash  Fork  Little 
Laramie  River  (7.5  miles  [ca.  12.1  km]  NW  of  Centennial 
[2,990  m])  [July]  (1;  DHKa),  Snowy  Range  Pass  ([3,200  m]) 
[June,  Aug.]  (33:  RTBe,  UASM);  Carbon  County,  Elk  Moun- 
tain (35.0  miles  [ca.  56.3  km]  NE  of  Saratoga  [3,400  m])  [July] 
(32;  USNM),  Hidden  Treasure  Gulch  (11.5  miles  [ca.  18.5  km] 
wsw  of  Encampment  [2,870  m])  [July]  (2;  DHKa),  Lake  Marie 
([3,230  m]  and  8  miles  [ca.  13  km]  sw  [2,740  m])  [July]  (7; 
DHKa),  North  French  Creek  (4  miles  [ca.  6.4  km]  w  of  Mirror 
Lake  [3,050  m])  [July]  (3;  DHKa),  Silver  Lake  ([3,170  m]) 
[July]  (32;  USNM). 

Doubtful  Records 

UNITED  STATES  OF  AMERICA — MASSACHUSETTS:  (1: 
UMRM). 

Nebria  acuta  quileute,  new  subspecies 

(Figures  16,  38,  46,  55) 

Nebria  acuta  LINDROTH  1961:71  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wash.,  Olym- 
pic N.  P.,  Olympic  Mts..  Boulder  Cr.  at  Olympic  Hot  Springs, 
2000',  20July71  DHKavanaugh  &  EAMartinko"/  "D.  H.  Ka- 
vanaugh  Collection"  [orange  label]/  "Holotype  Nebria  acuta 
quileute  Kavanaugh  del.  D.  H.  Kavanaugh  1976"  [red  label]/ 
"California  Academy  of  Sciences  Type  No.  12494."  ALLO- 
TYPE  (same  data  as  holotype)  also  in  CAS.  In  total,  135  PARA- 
TYPES  (81  males  and  54  females)  are  deposited  in  the  following 
collections:  CAS,  CNC,  DHKa,  LACM,  MCZ,  UASM, 
UIMI,  USNM,  UWBM. 

TYPE-LOCALITY. — Boulder  Creek  (at  Olympic  Hot  Springs). 
Olympic  National  Park,  Washington. 

DIAGNOSTIC  COMBINATION. — Pronotum  (Fig. 
16)  moderately  cordate,  basal  sinuation  of  lateral 
margin  moderately  short  and  deep,  basal  angle 
rectangular  or  slightly  acute,  midlateral  seta  ab- 
sent; elytra  slightly  shiny,  microsculpture  mod- 
erately impressed,  meshes  isodiametric  (broken) 
or  slightly  transverse,  elytral  silhouette  subrect- 
angular,  not  narrowed  basally,  humeral  angle 
not  rounded  or  obtuse,  humeral  carina  markedly 


KAVANAUGH:   NEW  NEARCTIC  NEBRIA 


99 


developed  and  projected  anterolaterally;  hind 
coxa  bi-  or  plurisetose  basally:  third  to  fifth  vis- 
ible abdominal  sterna  each  with  two  or  more 
pairs  of  posterior  paramedial  setae;  male  with 
apex  of  median  lobe  as  in  Figure  38;  female 
without  anterodorsal  sclerite  in  bursa  copulatrix 
(Fig.  46);  specimen  from  locality  on  Olympic 
Peninsula,  Washington  (Fig.  55). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  people  native  to  the  area 
including  the  type-locality,  the  Quileute  tribe. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  55; 
restricted  to  the  Olympic  Peninsula,  Washing- 
ton. I  have  studied  specimens  from  the  following 
localities. 

United  States  of  America 

WASHINGTON:  Olympic  National  Park  [Aug.]  (5;  LACM, 
MCZ),  Dosewallips  River  (at  Mascott  Campground)  [July]  (10; 
UWBM),  Eagle  Point  ([1,830  m])  [July]  (1;  CNC),  Olympic 
Hot  Springs  (Boulder  Creek  [610  m-780  m|)  [May-Aug.]  ( 120: 
CAS.  CNC,  DHKa,  UIMI,  USNM,  UWBM),  Sol  Due  Hot 
Springs  [June]  (1;  CAS). 

Nebria  sahlbergii  modoc,  new  subspecies 

(Figures  17,  56) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Calif.,  Modoc 
Co.,  Warner  Mts.,  4  mi.  e.  New  Pine  Creek,  Pine  Cr.,  5700', 
22May71  DHKavanaugh  &  EAMartinko"/  "D.  H.  Kavanaugh 
Collection"  [orange  label]/  "Holotype  Nebria  sahlbergii  mo- 
doc Kavanaugh  del.  D.  H.  Kavanaugh  1976"  [red  label]/ 
"California  Academy  of  Sciences  Type  No.  12513."  ALLO- 
TYPE  (same  data  as  holotype)  also  in  CAS.  In  total,  15  PARA- 
TYPES  (10  males  and  5  females)  are  deposited  in  the  following 
collections:  CAS,  DHKa,  UASM,  USNM. 

TYPE-LOCALITY. — Pine  Creek  (4  miles  [ca.  6.4  km]  E  of 
New  Pine  Creek),  Modoc  County,  California. 

DIAGNOSTIC  COMBINATION. — Pronotum  (Fig. 
17)  size  moderate,  basal  sinuation  of  lateral  mar- 
gin long  and  deep,  basal  angles  rectangular,  not 
denticulate,  lateral  explanation  broad  through- 
out its  length,  midlateral  seta  absent;  proepister- 
num  faintly  punctulate  anteriorly;  elytra  slightly 
short  and  narrowed  basally,  elytral  silhouette 
subrectangular,  humeral  angle  not  markedly 
rounded  or  obtuse,  humeral  carina  moderately 
developed,  slightly  projected  anterolaterally; 
specimen  from  locality  in  Warner  Mountains 
(northeastern  California  or  southcentral  Oregon) 
(Fig.  56). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  county  in  which  the  type- 
locality  is  found,  as  well  as  for  the  people  native 
to  the  region,  the  Modoc  tribe. 

GEOGRAPHICAL   DISTRIBUTION. — Figure  56; 


restricted  to  the  Warner  Mountains  of  north- 
eastern California  and  southcentral  Oregon.  I 
have  studied  specimens  from  the  following  lo- 
calities. 

United  States  of  America 

CALIFORNIA:  Modoc  County.  Cedar  Pass  [June]  (6; 
USNM),  New  Pine  Creek  (4  miles  [ca.  6.4  km]  E  at  Pine  Creek 
[1,740m])  [May]  (11:  DHKa). 

Nebria  sahlbergii  triad,  new  subspecies 

(Figures  18,  56) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Calif.,  Trin- 
ity Co.,  Trinity  Alps,  S.  F.  Salmon  R.  at  Big  Flat  Cmpgd., 
1490m,  18July75  D.H.  &  B.A.  Kavanaugh  Stop  #75-36"/  "D. 
H.  Kavanaugh  Collection"  [orange  label]/  "Holotype  Nebria 
sahlbergii  triad  Kavanaugh  del.  D.  H.  Kavanaugh  1976"  [red 
label]/  "California  Academy  of  Sciences  Type  No.  12514." 
ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In  total,  212 
PARATYPES  (98  males  and  114  females)  are  deposited  in  the 
following  collections:  CAS,  DHKa,  FMNH,  MCZ,  PUCA, 
UASM,  USNM. 

TYPE-LOCALITY. — South  Fork  Salmon  River  (at  Big  Flat 
Campground),  Trinity  County,  California. 

DIAGNOSTIC  COMBINATION. — Pronotum  (Fig. 
18)  large,  broad,  basal  sinuation  of  lateral  margin 
moderate  in  length  and  depth,  basal  angles  rect- 
angular, not  denticulate,  lateral  explanation 
broad  throughout  its  length,  midlateral  seta  ab- 
sent; proepisternum  smooth  anteriorly;  elytra 
moderate  in  length  and  basal  width,  elytral  sil- 
houette subrectangular,  humeral  angle  not 
markedly  rounded  or  obtuse,  humeral  carina 
moderately  developed,  slightly  projected  an- 
terolaterally; specimen  from  locality  in  Klamath 
Mountains  system  of  the  Coast  Ranges  (north- 
western California  or  southwestern  Oregon) 
(Fig.  56). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Trinity  Alps,  in  which  the 
type-locality  is  found.  I  use  a  synonym  of  'trin- 
ity' for  this  purpose. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  56; 
restricted  to  the  Klamath  Mountains  system  of 
northwestern  California,  probably  also  in  south- 
western Oregon.  I  have  studied  specimens  from 
the  following  localities. 

United  States  of  America 

CALIFORNIA:  Siskiyou  County,  Gulick  Creek  (at  road  to 
Josephine  Lake  [1,550  m])  [Aug.]  (1:  CAS):  Trinity  County, 
Boulder  Creek  (at  Goldfield  Campground  [1,070  m])  [July]  (56; 
CAS,  DHKa).  Coffee  Creek  (at  Coffee  Creek  Ranch  [1,070 
m])  [July]  (32;  CAS,  DHKa),  Morris  Meadows  ([1,340  m]) 
[Aug.]  (1:  PUCA),  South  Fork  Salmon  River  (at  Big  Flat 
Campground  [1,490  m])  [June-Aug.]  (124:  CAS,  DHKa, 
FMNH,  MCZ.  UASM,  USNM). 


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Nebria  lituyae,  new  species 

(Figures  29,  57) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "4  June  1977  3356' 
under  stones,  summit  Mt.  Blunt,  2  mi.  s.  Lituya  Bay,  SE 
Alaska  D.  Mann  coll.  102"/  "Holotype  Nebria  lituyae  Kavan- 
augh  det.  D.  H.  Kavanaugh  1979"  [red  label]/  "California 
Academy  of  Sciences  Type  No.  13460."  ALLOTYPE  (same 
data  as  holotype)  also  in  CAS.  In  total,  21  PARATYPES  (11 
males  and  10  females)  are  deposited  in  the  following  collec- 
tions: CAS,  DHKa,  DMan,  UASM,  USNM. 

TYPE-LOCALITY. — Mt.  Blunt,  2  miles  [ca.  3.2  km]  s  of  Li- 
tuya Bay,  Alaska. 

DIAGNOSTIC  COMBINATION. — Head  width 
moderate  or  slightly  broadened,  eyes  moderate- 
ly convex,  each  epilobe  of  mentum  with  apical 
tooth  short  or  moderately  long;  pronotum  with 
lateral  explanation  broad  throughout  its  length, 
basal  angles  rectangular,  basal  sinuation  of  lat- 
eral margin  moderate  in  length  and  depth,  mid- 
lateral  seta  absent;  prosternal  intercoxal  process 
(Fig.  29)  lanceolate,  completely  margined  (even 
at  apex);  elytra  slightly  elongate,  narrow,  with 
or  without  faint  metallic  violet  reflection,  elytral 
silhouette  subovoid,  slightly  to  moderately  nar- 
rowed basally,  humeral  angle  not  markedly 
rounded  or  obtuse,  humeral  carina  slightly  de- 
veloped and  projected  anterolaterally;  hind  coxa 
bi-  or  plurisetose  basally;  third  to  fifth  visible 
abdominal  sterna  each  with  two  or  more  pairs 
of  posterior  paramedial  setae;  specimen  from  lo- 
cality in  southeastern  Alaska  (mainland)  (Fig. 
57). 

DERIVATION  OF  TAXON  NAME. — This  species 
is  named  for  Lituya  Bay,  near  which  the  type- 
locality  is  found. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  57; 
known  only  from  southeastern  Alaska,  near  Li- 
tuya Bay.  I  have  studied  specimens  from  the 
following  locality. 

United  States  of  America 

ALASKA:  Mount  Blunt  (2  miles  [ca.  3.2  km]  s  of  Lituya  Bay 
[720  m-760  m])  [June]  (23;  CAS,  DHKa,  DMan,  UASM, 

USNM). 

Nebria  arkansana  edwardsi,  new  subspecies 

(Figures  30,  33,  39,  47,  58) 

Nebria  sahlhergi;  auctorum.- — EDWARDS   1975:48.  • — MANK 

1934:74. 
Nebria  arkansana;  LINDROTH  1961:70  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Mont.,  Glacier 
N.  P.,  Livingston  Range,  Logan  Pass  Area,  ne.  slope  Mt. 
Clements,  7100',  17July72  DHKavanaugh  &  JGEdwards"/ 
"D.  H.  Kavanaugh  Collection"  [orange  label]/  "Holotype 
Nebria  arkansana  edwardsi  Kavanaugh  det.  D.  H.  Kavanaugh 


1976"  [red  label]/  "California  Academy  of  Sciences  Type  No. 
12495."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In 
total,  1872  PARATYPES  (1060  males  and  812  females)  are  de- 
posited in  the  following  collections:  AMNH,  ANSP,  BFCa, 
CAS,  CNC,  CUIC,  DBUM,  DHKa,  DJLa,  FMNH,  INHS, 
JSch,  JVMa,  KUSM,  LRus,  MCZ,  MSU,  OSUO,  PMNH, 
PURC,  ROM,  RTBe,  SJSC,  UAFA,  UASM,  UBC,  UIMI, 
USNM,  UWBM. 

TYPE-LOCALITY. — Logan  Pass,  Glacier  National  Park, 
Montana. 


NOTES  ON  NOMENCLATURE  AND  TYPES. — 
Lindroth  (1961:70)  designated  the  valley  of  the 
Upper  San  Juan  River,  Archuleta  County,  Col- 
orado, as  type-locality  for  N.  arkansana  Casey. 
As  a  result,  the  nominal  subspecies  of  N.  ar- 
kansana is  one  restricted  to  the  southern  Rocky 
Mountains  (southcentral  Wyoming  and  further 
south).  The  name  presented  here  is  for  the 
widely  distributed  subspecies  left  unnamed  by 
Lindroth's  choice  of  type-locality. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  relative  width;  pronotum  with  lateral 
explanation  broad  throughout  its  length,  basal 
angles  rectangular  or  slightly  acute,  basal  sin- 
uation of  lateral  margin  moderately  deep,  mid- 
lateral  seta  absent;  elytra  slightly  to  moderately 
dull,  microsculpture  with  meshes  isodiametric 
or  slightly  transverse,  elytral  silhouette  rectan- 
gular or  subrectangular,  broad  basally,  basal 
margination  (Fig.  30)  straight  or  slightly  con- 
cave, merged  smoothly  with  lateral  margination, 
humeral  angle  not  markedly  rounded  or  obtuse, 
humeral  carina  absent  or  very  slightly  devel- 
oped; hind  wing  with  full  reflexed  apex  distal  to 
stigma  (Fig.  33);  hind  coxa  bi-  or  plurisetose  ba- 
sally; third  to  fifth  visible  abdominal  sterna  each 
with  two  or  more  pairs  of  posterior  paramedial 
setae;  male  with  median  lobe  (Fig.  39)  broad, 
with  apex  short  and  narrow;  female  with  pos- 
terodorsal  sclerite  of  bursa  copulatrix  as  in  Fig- 
ure 47;  specimen  from  locality  in  Rocky  Moun- 
tain region,  but  only  north  or  west  of  Utah  and 
southern  Wyoming  (Fig.  58). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  subspecies  in  honor  of 
J.  Gordon  Edwards,  whose  keen  interest  in  life 
at  high  altitude  has  fostered  my  own. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  58; 
widespread,  distributed  in  Rocky  Mountain  re- 
gion from  southern  Yukon  Territory  south  to 
northeastern  Nevada,  Idaho,  and  northern  Wy- 
oming. I  have  studied  specimens  from  the  fol- 
lowing localities. 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


101 


Canada 

ALBERTA:  Banff  National  Park,  Banff  [Aug. -Sep.)  (3; 
CUIC,  UASM),  Boom  Lake  ([1.830  m|)  [July]  (1;  CNC),  Bow 
Lake  [Aug.]  (1;  UASM),  Bow  River  ([1,830  m])  [Aug.]  (6; 
CNC),  Johnson  Creek  Trail  (at  Johnson  Creek  Campground) 
[July]  (1;  ROM),  Lake  Agnes  [Aug.]  (1:  PMNH),  Lake  Louise 
[June-Sep.]  (16;  AMNH,  CAS,  CNC,  MCZ,  UBC),  Ptarmigan 
Mountain  ([2,590  m])  [Aug.]  (13;  CNC),  Spray  Lake  [July]  (1; 
JVMa),  Sunset  Pass  ([1,950  m])  [Aug.]  (4;  CNC);  Jasper  Na- 
tional Park  [Aug.]  (2;  AMNH),  Amethyst  Lake  (and  Ame- 
thyst Lake  Trail  [1,710  m])  [July]  (10;  DHKa,  UASM),  Angel 
Glacier  [Sep.]  (1;  RTBe),  Hardisty  Creek  (1  mile  [ca.  1.6  km] 
s  on  Highway  93)  [July]  (1;  RFre),  Jasper  [Aug.]  (3:  USNM), 
Miette  Hot  Springs  [Aug.]  (1;  RTBe),  Miette  River  (at  Mead- 
ow Creed  [1,100  m])  [July-Aug.]  (22;  DHKa,  UASM),  Mount 
Edith  Cavell  ([1,360  m]  and  below  Angel  Glacier)  [July]  (3: 
DHKa,  UAFA),  Portal  Creek  Trail  ([1.950  m])  [July]  (3; 
USNM);  Waterton  Lakes  National  Park  [July-Aug.]  (5; 
UASM,  UWBM),  Cameron  Lake  (creek  below  lake)  [July] 
(7;  USNM),  Red  Rock  Canyon  [July]  (2:  CAS),  Waterton  Park 
(Cameron  Creek)  [July]  (36;  CNC,  MCZ,  UASM);  Other  lo- 
calities, Bellevue  (2  miles  [ca.  3.2  km]  w  at  Gold  Creek  [1,370 
m])  [July]  (1;  DHKa),  Blackstone  Gap  [Aug.]  (2;  CNC).  Blair- 
more  (1;  USNM),  Blakiston  Brook  [July]  (10;  UASM),  Car- 
bondale  River  (near  Lost  Creek)  [Sep.|  (1;  DJLa),  Cline  River 
(at  Highway  11)  [Aug.]  (2:  DHKa),  Gap  [Aug.]  (1;  CAS), 
Gorge  Creek  [July]  (2;  UASM),  Happy  Valley  (Pincher  Creek) 
[Aug.]  (11;  CAS,  UASM),  Highwood  Pass  ([2,380  m])  [July] 
(15;  CNC),  Highwood  River  [Aug.]  (1;  CAS),  Marmot  Creek 
(10  miles  [ca.  16  km]  sw  of  Kananaskis  Forest  Experiment 
Station  [1.520  m])  [Aug.]  (10:  CNC),  Mill  Creek  Ranger  Sta- 
tion [Aug.]  (2;  CAS),  Pincher  Creek  ([1,620  m])  [July]  (3; 
DJLa),  Ram  River  area  [July]  (2;  BFCa),  Sheep  River  (4.5 
miles  [ca.  7.2  km]  above  Gorge  Creek)  [Aug.]  (7:  DHKa). 
West  Branch  Castle  River  ([1,340  m])  [Aug.]  (6:  DJLa).  BRIT- 
ISH COLUMBIA:  Glacier  National  Park,  Asulkan  Glacier  area 
[Aug.]  (I;  CNC),  Glacier  [July-Aug.]  (6:  CAS,  CNC,  PURC), 
Rogers  Pass  ([1,370  m])  [Aug.]  (2;  ANSP,  CAS);  Kootenay 
National  Park  [Aug.]  (3;  CNC),  Kimpton  Creek  ([1,220  m]) 
[July]  (2;  CNC),  Kindersley  Pass  ([2,130  m])  [July]  (4;  CNC), 
Kootenay  River  (at  Highway  IB)  [Aug.]  (1:  UASM),  Marble 
Canyon  [July]  (1;  CNC),  Sinclair  Creek  ([1,830  m]  and  at 
Highway  13)  [June-Aug.]  (25;  CAS,  CNC,  UASM,  UBC), 
Vermillion  Creek  [July]  (4;  CAS,  MSU,  UBC);  Yoho  National 
Park  [July]  (3;  CAS),  Amiskwi  River  (and  at  Amiskwi  Falls 
[1,830  m])  [Aug.]  (34;  CNC),  Field  (and  3  miles  [ca.  4.8  km] 
E  on  Kicking  Horse  River)  [June,  Aug.]  (4:  CAS,  USNM). 
Lake  Oesa  ([2,130  m])  [July]  (18;  CNC),  Lake  O'Hara  [July] 
(10;  CAS,  CNC,  UBC),  Otterhead  River  ([1,680  m])  [Aug.- 
Sep.]  (29;  CNC,  DHKa),  Yoho  River  ([  1 ,400  m],  4  to  12  miles 
[ca.  6.4toca.  19km]  E  of  Field,  at  Takakkaw  Falls  [1,520m]) 
[July-Aug.]  (25;  CNC,  DHKa,  UASM);  Other  localities,  Bo- 
sewell  [Aug.]  (1;  UASM),  Cabin  Creek  (NE  of  Fernie,  near 
Michel)  [July]  (13;  CNC,  UASM),  Creston  (8  miles  [ca.  13 
km]  w)  [June]  (3;  CNC),  Crowsnest  Pass  (2  miles  [ca.  3.2  km] 
w  [1,280  m]  and  5  miles  [ca.  8  km]  w  on  Elk  River  [1,220  m]) 
[July]  (51:  DHKa),  Edgewood  [July-Aug.)  (3;  CAS),  Elk 
Creek  (E  of  Fernie)  [July]  (4;  UASM),  Elk  River  (10.7  miles 
[ca.  17.2  km]  w  of  Fernie)  [July]  (3:  ROM),  Erickson  (Arrow 
Creek)  [Aug.]  (12:  CAS,  UBC),  Fernie  (Coal  Creek  [1,680  m]. 
Fairy  Creek.  Lizard  Creek)  [July-Aug.)  (63:  BFCa.  CAS, 
CNC,  DBUM,  FMNH,  MSU,  UBC),  Flathead  [June]  (1; 
CUIC),  Fort  Steele  [June,  Sep.]  (2;  CAS,  UBC),  Fortysixmile 
Camp  (at  Hope  Trail)  [July]  (4;  CAS,  CNC,  UBC),  Hope 


Mountains  [July-Sep.]  (6;  CNC,  INHS,  MCZ),  Hosmer  (Elk 
River)  [July]  (1;  CAS),  Lorna  [June]  (2;  CAS,  UBC),  Manning 
Provincial  Park  (Nicomen  Ridge)  [July]  (10;  CAS,  CNC, 
MSU,  UBC),  Michel  (5;  UASM),  Mount  Copeland  ([1,980  m]) 
[Aug.]  (21;  CNC),  Radium  Hot  Springs  [Aug.]  (6;  CAS,  CNC, 
UBC),  Slocan  [Aug.]  (2;  BFCa),  South  Fork  Creek  [Aug.]  (1; 
UWBM),  Summit  Creek  (near  Creston)  [Aug.]  (5;  UASM), 
Tetsa  River  (Alaska  Highway  mile  378  and  mile  390)  [Aug.] 
(3;  DHKa,  MSU),  Vernon  [July]  (1;  CAS),  Whipsaw  Creek 
(at  Hope  Trail  [1,460  m])  [July]  (1;  UBC),  Wyndel  [Sep.]  (1; 
CNC).  YUKON  TERRITORY:  Rancheria  (Swift  River)  [Aug.]  (6; 
CNC,  UASM). 

United  States  of  America 

IDAHO:  Blaine  County,  Alturas  Lake  (at  Alturas  Inlet 
Campground  [2,130  m])  [Aug.]  (11;  DHKa),  Cherry  Creek  (0. 1 
miles  [ca.  0.2  km]  NW  of  Highway  93  [2,230  m])  [Aug.]  (6: 
DHKa),  Dollarhide  Summit  (2  miles  [ca.  3.2  km]  E  at  North 
Fork  Warm  Springs  Creek  [2,350  m-2,410  m])  [Aug.]  (9: 
DHKa),  Galena  [July]  (1:  BFCa),  Galena  Summit  (2.4  miles 
[ca.  3.9  km]  NE  [2,320  m])  [Aug.]  (12;  DHKa),  Hyndman 
Creek  [July]  (22:  UWBM),  Ketchum  (2  miles  [ca.  3.2  km] 
NNW  at  Big  Wood  Creek  [1,830  m])  [Aug.]  (28;  DHKa),  North 
Fork  Warm  Springs  Creek  (10  miles  [ca.  16  km]  w  of  Ketchum 
[1,890  m])  [Aug.]  (92;  DHKa),  Sun  Valley  [Sep.]  (2;  JSch); 
Camas  County,  Carrie  Creek  (32  miles  [ca.  51  km]  [2,440  m] 
and  36  miles  [ca.  58  km]  [2,100  m]  ESE  of  Ketchum)  [Aug.] 
(106;  DHKa),  South  Fork  Boise  River  (14  miles  [ca.  23  km) 
E  of  Featherville  at  Skeleton  Creek  [1,550  m],  34  miles  [ca.  55 
km]  E  of  Featherville  at  Bear  Creek  [1,830  m])  [Aug.]  (157: 
DHKa):  Cassia  County,  Rock  Creek  (15  miles  [ca.  24  km]  s 
of  Hansen,  at  Rock  Creek  Ranger  Station)  [June-July]  (6; 
UIMI);  Custer  County,  Bayhorse  Creek  (7  miles  [ca.  11  km) 
NW  of  Highway  93  [2,440  m])  [Aug.]  (3;  DHKa),  Fourth  of 
July  Creek  (3  miles  [ca.  4.8  km]  E  of  Highway  93  [2,230  m]) 
[Aug.]  (58:  DHKa).  Stanley  [July]  (1;  UWBM).  Trail  Creek 
Summit  (2  miles  [ca.  3.2  km]  NE  at  Summit  Creek  [2,320  m]) 
[Aug.]  (9;  DHKa);  Fremont  County,  Island  Park  [July]  (3: 
BFCa);  Shoshone  County,  Avery  (7  miles  [ca.  1 1  km]  E)  [July] 
(1;  UIMI);  Valley  County,  Dagger  Falls  [July]  (2;  BFCa), 
Middle  Fork  Salmon  River  (near  Boundary  Creek)  [Aug.]  (7; 
RTBe);  County  unknown,  Twin  Creek  Forest  Camp  ([1.520 
m])  [July]  (57:  UWBM).  MONTANA:  Carbon  County,  Quad 
Creek  (17  miles  [ca.  27  km]  sw  of  Red  Lodge  [3,050  m])  [July) 
(66;  DHKa);  Fergus  County,  East  Fork  Rock  Creek  ([1,710 
m))  [July]  (42;  DHKa):  Flathead  County,  Marias  Pass  (10 
miles  [ca.  16  km]  w  of  summit  [1,370  m])  [July]  (11:  LRus). 
South  Fork  Flathead  River  (at  Hungry  Horse  Reservoir  [1,220 
m])  [July]  (3:  LRus);  Gallatin  County,  Fairy  Lakes  [Aug.]  (I; 
CUIC);  Glacier  National  Park  [July-Aug.]  (6;  CUIC),  Baring 
Creek  (at  St.  Mary  Lake)  [Aug.]  (3:  UWBM),  Grinnel  Falls 
([1,860  m])  [Aug.]  (1;  SJSC),  Grinnel  Glacier  ([1,860  m-1,890 
m])  [Aug.]  (21:  DHKa,  SJSC,  USNM),  Grinnel  Lake  (at  inlet 
[1,540m])  [Aug.](l;  DHKa).  Lake  McDonald  (and  at  Sprague 
Creek  Campground)  [July-Aug.]  (4;  CAS,  RTBe),  Lake  Sher- 
burne  (at  Canyon  Creek)  [July]  (1:  CAS),  Logan  Pass  ([2,030 
m-2,130  m]  and  5  miles  [ca.  8  km]  w  [1,770  m])  [July-Sep.] 
( 16;  DHKa,  JSch,  RTBe,  UASM),  Morning  Eagle  Falls  ([1,680 
m])  [July]  (1:  SJSC),  Mount  Clements  (NE  slope  [2,160  m]) 
[July]  (5;  DHKa),  Piegan  Pass  ([2,410  m])  [Aug.)  (I;  SJSC), 
St.  Mary  [Aug.]  (8;  SJSC,  UASM):  Madison  County,  Beaver 
Creek  ([1,920  m])  [Aug.]  (1:  KUSM),  Ennis  [July)  (2;  UIMI); 
Mineral  County,  Haugan  [Aug.)  (14;  KUSM),  Lookout  Pass 
([1,280  m))  [July]  (1;  LRus),  Saltese  [July,  Sep.)  (3:  MSU, 


102 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


SJSC):  Sweetgrass  County,  Big  Timber  Creek  (at  Half  Moon 
Campground  [2,230  m-2,290  m])  [July]  (1;  DHKa).  NEVADA: 
Elko  County,  Lamoille  Creek  (at  Lower  Lamoille  Camp- 
ground [1,920  m])  [Aug.]  (35;  DHKa),  Thomas  Creek  (at 
Thomas  Creek  Campground  [2,320  m-2,380  m])  [Aug.]  (130; 
DHKa).  OREGON:  Baker  County,  Cornucopia  [June-July]  (35; 
BFCa,  UWBM);  Union  County,  Elgin  (and  9  miles  [ca.  14 
km]  NW  at  Little  Phillips  Creek  [1,300  m])  [May,  Aug.]  (6; 
DHKa),  Langdon  Lake  area  (SE  of  summit)  [Aug.]  (2; 
UWBM),  Tollgate-Elgin  Road  [June]  (3;  UWBM);  Wallowa 
County  [Aug.]  (1;  JSch),  Aneroid  Lake  ([2,320  m])  [July]  (2; 
FMNH),  East  Fork  Eagle  Creek  [July]  (9;  UWBM),  Indian 
Crossing  (Imnaha  River)  [Aug.]  (2;  MSU,  OSUO),  Lostine 
River  ([1,310  m],  at  French  Campground,  at  Two  Pan  Camp- 
ground [1,710  m- 1,980  m])  [Aug.]  (112;  DHKa,  USNM, 
UWBM),  Lost  Lake  [Aug.]  (1;  UWBM),  Wallowa  Lake  [July- 
Aug.]  (10;  DHKa,  UWBM),  Wallowa  River  (above  inlet  to 
Wallowa  Lake)  [May]  (1;  DHKa).  WASHINGTON:  Walla  Walla 
County,  Walla  Walla  [Aug.]  (1;  UWBM).  WYOMING:  Big 
Horn  County,  Granite  Pass  (8  miles  [ca.  13  km]  sw  on  Granite 
Creek  [2,380  m])  [July]  (5;  DHKa),  Shell  Creek  (11  miles  [ca. 
18  km]  sw  of  Granite  Pass  [2,130  m])  [July]  (14;  DHKa),  West 
Tensleep  Creek  ([3,140  m-3,200  m])  [July]  (17;  DHKa);  John- 
son County,  Clear  Creek  (12  miles  [ca.  19  km]  w  of  Buffalo 
[2,190  m])  [July]  (1;  DHKa),  South  Fork  Clear  Creek  (16  miles 
[ca.  26  km]  w  of  Buffalo  [2,350  m],  at  South  Fork  Campground 
[2,350  m])  [July-Aug.]  (6;  DHKa,  FMNH);  Park  County,  Fox 
Creek  (at  Highway  212)  [Aug.]  (1;  CAS);  Sheridan  County, 
Little  Tongue  River  (13  miles  [ca.  21  km]  wsw  of  Dayton 
[2,380  m])  [July]  (22;  DHKa);  Washakie  County,  Tensleep  (6 
miles  [ca.  10  km]  [1,550  m]  and  11  miles  [ca.  18  km]  [1,890  m] 
NE  on  Tensleep  Creek)  [July]  (4;  DHKa):  Yellowstone  Na- 
tional Park  [Aug.]  (25;  USNM);  County  unknown,  Bighorn 
Reserved;  MCZ). 

Nebria  arkansana  oowah,  new  subspecies 

(Figures  40,  48,  58) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Utah,  Grand 
Co.,  LaSal  Mts.,  Mill  Cr.  at  Oowah  Lk.,  8800',  2August71 
DHKavanaugh  &  EAMartinko"/  "D.  H.  Kavanaugh  Collec- 
tion" [orange  label]/  "Holotype  Nebria  arkansana  oowah  Ka- 
vanaugh del.  D.  H.  Kavanaugh  1976"  [red  label]/  "California 
Academy  of  Sciences  Type  No.  124%."  ALLOTYPE  (same 
data  as  holotype)  also  in  CAS.  In  total,  75  PARATYPES  (54 
males  and  21  females)  are  deposited  in  the  following  collec- 
tions: CAS,  DHKa,  MCZ,  UASM,  USNM. 

TYPE-LOCALITY. — Mill  Creek  (at  Oowah  Lake),  Grand 
County,  Utah. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  relative  width;  pronotum  with  lateral 
explanation  broad  throughout  its  length,  basal 
angles  rectangular  or  slightly  acute,  basal  sin- 
uation  of  lateral  margin  moderately  deep,  mid- 
lateral  seta  absent;  elytra  with  humeral  angle  not 
markedly  rounded  or  obtuse,  humeral  carina 
moderately  developed,  elytral  silhouette  sub- 
rectangular,  broad  basally,  hindwing  full-sized; 
hind  coxa  bi-  or  plurisetose  basally;  third  to  fifth 
visible  abdominal  sterna  each  with  two  or  more 
pairs  of  posterior  paramedial  setae;  male  with 


median  lobe  broad,  apex  as  in  Figure  40;  female 
with  posterodorsal  sclerite  of  bursa  copulatrix 
as  in  Figure  48;  specimen  from  locality  in  La  Sal 
Mountains  (eastern  Utah)  (Fig.  58). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  small,  sheltered  lake  at  the 
type-locality. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  58; 
restricted  to  the  La  Sal  Mountains  of  eastern 
Utah.  I  have  studied  specimens  from  the  follow- 
ing localities. 

United  States  of  America 

UTAH:  Grand  County,  Mill  Creek  (at  Oowah  Lake  [2,680 
m])  [Aug.]  (74;  CAS,  DHKa,  MCZ,  USNM);  San  Juan  Coun- 
ty, Mill  Creek  Valley  ([2,900  m])  [Aug.]  (3;  UASM). 

Nebria  arkansana  uinta,  new  subspecies 

(Figures  35,  41,  58) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Utah,  Summit 
Co.,  Uinta  Mts.,  Hwy.  150,  Lost  Lk.,  9800',  14Aug71  DHKa- 
vanaugh"/ "D.  H.  Kavanaugh  Collection"  [orange  label]/ 
"Holotype  Nebria  arkansana  uinta  Kavanaugh  det.  D.  H. 
Kavanaugh  1976"  [red  label]/  "California  Academy  of  Sci- 
ences Type  No.  12497."  ALLOTYPE  (same  data  as  holotype) 
also  in  CAS.  In  total,  93  PARATYPES  (44  males  and  49  females) 
are  deposited  in  the  following  collections:  CAS.  DHKa,  MCZ, 
UASM,  USNM. 

TYPE-LOCALITY. — Lost  Lake,  Summit  County,  Utah. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  relative  width;  pronotum  with  lateral 
explanation  broad  throughout  its  length,  basal 
angles  rectangular  or  slightly  acute,  basal  sin- 
uation  of  lateral  margin  moderately  deep,  mid- 
lateral  seta  absent;  elytra  with  humeral  angle  not 
markedly  rounded  or  obtuse,  humeral  carina 
moderately  developed;  elytral  silhouette  sub- 
rectangular  or  subovoid,  slightly  narrowed  ba- 
sally; hindwing  reduced  in  length  and  width  (Fig. 
35);  hind  coxa  bi-  or  plurisetose  basally;  third  to 
fifth  visible  abdominal  sterna  each  with  two  or 
more  pairs  of  posterior  paramedial  setae;  male 
with  median  lobe  as  in  Figure  41 ;  specimen  from 
locality  in  Uinta  Mountains  or  in  Wasatch 
Mountains  at  or  north  of  Weber  River,  Utah 
(Fig.  58). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Uinta  Mountains,  in  which 
the  type-locality  is  found. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  58; 
restricted  to  the  Uinta  Mountains  and  the  Wa- 
satch Mountains  at  or  north  of  Weber  River 
(northern  Utah).  I  have  studied  specimens  from 
the  following  localities. 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


103 


United  States  of  America 

UTAH:  Cache  County,  Logan  River  (13. 7  miles  [ca.  22  km] 
E  of  Logan  at  Cottonwood  Creek  [1,740  m])  [Aug.]  (65;  CAS, 
DHKa,  MCZ,  UASM):  Duchesne  County,  Painter  Basin 
([3,470  m])  [Sep.]  (1;  USNM):  Summit  County,  Lost  Lake 
([2,990  m])  [Aug.]  (7;  DHKa),  Upper  Provo  River  (20.8  miles 
[ca.  33  km]  E  of  Kamas  [2,600  m],  at  Highway  150)  [Aug.]  (6: 
DHKa,  USNM);  Uintah  County,  Big  Brush  Creek  (7  miles 
[ca.  11  km]  WNW  of  Highway  44  [2,620  m])  [Aug.]  (14; 
DHKa):  Weber  County,  Ogden  (6  miles  [ca.  10  km]  E  on  Og- 
den  River  [1,490  m])  [Aug.]  (2;  DHKa). 

Nebria  fragilis  teewinot,  new  subspecies 

(Figures  35,  42,  59) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wyo.,  Teton 
N.  P.,  Teton  Mts.,  Mt.  Teewinot,  stream  on  se.  slope,  7100'- 
9000',  3Aug.73  DHKavanaugh  Family"/  "D.  H.  Kavanaugh 
Collection"  [orange  label]/  "Holotype  Nebria  fragilis  teewinot 
Kavanaugh  del.  D.  H.  Kavanaugh  1976"  [red  label]/  "Cali- 
fornia Academy  of  Sciences  Type  No.  12500."  ALLOTYPE 
(same  data  as  holotype)  also  in  CAS.  In  total,  377  PARATYPES 
(219  males  and  158  females)  are  deposited  in  the  following 
collections:  CAS,  DHKa,  KUSM,  MCZ,  SJSC,  UASM, 
USNM. 

TYPE-LOCALITY. — Mount  Teewinot  (southeast  slope). 
Grand  Teton  National  Park,  Wyoming. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  relative  width;  pronotum  with  lateral 
explanation  broad  throughout  its  length,  basal 
angles  rectangular  or  slightly  acute,  basal  sin- 
uation  of  lateral  margin  moderately  deep,  mid- 
lateral  seta  absent;  elytra  slightly  shiny,  micro- 
sculpture  faintly  impressed,  with  meshes 
irregularly  isodiametric  or  slightly  transverse, 
with  or  without  faint  metallic  green  or  violet  re- 
flection, elytral  silhouette  subrectangular  or  sub- 
ovoid,  slightly  narrowed  basally,  humeral  angle 
not  markedly  rounded  or  obtuse,  humeral  carina 
moderately  developed;  hindwing  reduced  in 
length  and  width  (Fig.  35);  hind  coxa  bi-  or  plu- 
risetose  basally;  third  to  fifth  visible  abdominal 
sterna  each  with  two  or  more  pairs  of  posterior 
paramedial  setae;  male  with  median  lobe  as  in 
Figure  42;  specimen  from  locality  in  westcentral 
Wyoming  (Fig.  59). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  type-locality. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  59; 
restricted  to  mountain  ranges  of  the  Rocky 
Mountain  system  in  westcentral  Wyoming.  I 
have  studied  specimens  from  the  following  lo- 
calities. 

United  States  of  America 

WYOMING:  Grand  Teton  National  Park,  Mica  Lake  ([2,970 
m])  [July]  (1;  DHKa),  Mount  Teewinot  (SE  slope  [2,160  m- 
2,740  m])  [Aug.]  (79;  DHKa),  South  Fork  Cascade  Canyon 


([3,080  m])  [July]  (1:  SJSC),  South  Fork  Garnet  Canyon 
([2,820  m])  [July]  (1;  SJSC),  Timberline  Lake  area  [July]  (1; 
DHKa);  Lincoln  County,  White  Creek  (20  miles  [ca.  32  km] 
SE  of  Alpine  Junction  [1,950  m-2,190  m])  [Aug.]  (40;  DHKa), 
Wolf  Creek  (16.1  miles  [ca.  26  km]  sw  of  Hoback  Junction 
[1,770  m-1,780  m])  [July-Aug.]  (137:  CAS,  DHKa,  MCZ, 
UASM,  USNM);  Sublette  County,  Green  River  Lakes  (west 
shores  [2,440  m-2,590  m])  [July]  (53;  DHKa),  Hoback  River 
(14  miles  [ca.  23  km]  sw  of  Highway  187/189  [2,440  m])  [July] 
(11;  DHKa):  Teton  County,  Alaska  Basin  (near  Sunset  Lake 
[2,940  m])  [July]  (1;  SJSC),  Granite  Creek  (at  Granite  Falls 
[2,160  m])  [July]  (37;  DHKa),  Togwotee  Pass  ([2,960  m]  and 
3.7  miles  [ca.  6.0  km]  WNW  at  Blackrock  Creek  [2,770  m]) 
[Aug.]  (20:  DHKa,  KUSM). 

Nebria  zioni  oasis,  new  subspecies 

(Figures  5,  60) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Utah,  Wash- 
ington Co.,  Pine  Valley  Mts.,  Leeds  Cr.  at  Oak  Grove 
Cmpgd.,  6300' -6500',  20June71  DHKavanaugh  &  EAMartin- 
ko"/  "D.  H.  Kavanaugh  Collection"  [orange  label]/  "Holo- 
type Nebria  zioni  oasis  Kavanaugh  det.  D.  H.  Kavanaugh 
1976"  [red  label]/  "California  Academy  of  Sciences  Type  No. 
12518."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In 
total,  46  PARATYPES  (18  males  and  28  females)  are  deposited 
in  the  following  collections:  CAS,  DHKa,  UASM,  USNM. 

TYPE-LOCALITY. — Leeds  Creek  at  Oak  Grove  Camp- 
ground, Washington  County,  Utah. 

DIAGNOSTIC  COMBINATION. — Body  and  ap- 
pendages dark  (piceus  or  black);  antennal  scape 
(Fig.  5)  medium  length,  straight,  almost  cylin- 
drical (slightly  narrowed  basally);  each  epilobe 
of  mentum  with  tooth  moderately  long;  prono- 
tum with  lateral  explanation  moderately  broad 
throughout  its  length,  basal  angles  rectangular 
or  slightly  acute  and  projected  posteriorly,  lat- 
eral margin  without  subapical  sinuation  but  with 
basal  sinuation  moderately  long  and  deep,  mid- 
lateral  seta  absent;  elytra  not  vaulted,  slightly 
flattened,  elytral  silhouette  ovoid,  markedly  nar- 
rowed basally,  humeral  angle  markedly  rounded 
and  obtuse,  humeral  carina  markedly  devel- 
oped, fifth  and  seventh  elytral  intervals  without 
setiferous  pores;  hind  coxa  bi-  or  plurisetose 
basally;  third  to  fifth  visible  abdominal  sterna 
each  with  two  or  more  pairs  of  posterior  para- 
medial  setae;  specimen  from  locality  in  Pine 
Valley  Mountains  (Utah)  (Fig.  60). 

DERIVATION  OF  TAXON  NAME. — This  species 
is  named  for  the  oasislike  nature  of  the  type- 
locality — a  spot  of  green,  as  it  were,  in  an  oth- 
erwise dry  land. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  60; 
restricted  to  the  Pine  Valley  Mountains  of  south- 
western Utah.  I  have  studied  specimens  from 
the  following  locality. 


104 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


United  States  of  America 

UTAH:  Washington  County,  Leeds  Creek  (at  Oak  Grove 
Campground  [1-920  m-1,980  m])  [June]  (46;  CAS,  DHKa, 
UASM,  USNM). 

Nebria  obliqua  chuskae,  new  subspecies 

(Figures  19,  34,  61) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "USA  Arizona 
Apache  Co.,  sw.  Red  Rock  Chuska  Mts.  Wagon  Wheel  Camp 
Gr.  aspen-fir  forest  2260m.  Loc.  11  July  11-12,  1976' V 
"SOUTHWESTERN  USA  EXP.— 1976  G.E.Ball,  J.M. 
Campbell,  P.M.Hammond"/  "Holotype  Nebria  obliqua  chus- 
kae Kavanaugh  det.  D.  H.  Kavanaugh  1979"  [red  label]/ 
"California  Academy  of  Sciences  Type  No.  13461."  ALLO- 
TYPE  (same  data  as  holotype)  also  in  CAS.  In  total,  16  PARA- 
TYPES  (10  males  and  6  females)  are  deposited  in  the  following 
collections:  CAS,  MCZ,  UASM,  USNM. 

TYPE-LOCALITY. — Lukachukai  Creek  (at  Wagon  Wheel 
Campground),  Apache  County,  Arizona. 

DIAGNOSTIC  COMBINATION. — Mandible 
markedly  explanate  laterally,  broad;  tooth  of 
mentum  bifid,  with  notch  V-shaped;  pronotum 
(Fig.  19)  relatively  long  and  narrow,  moderately 
cordate,  moderately  narrowed  basally,  widest 
anterior  to  middle,  slightly  shiny  (microsculp- 
ture  faintly  or  moderately  impressed,  meshes 
isodiametric),  basal  margin  not  or  only  slightly 
sinuate  laterally,  lateral  bead  narrow,  obliterat- 
ed basally  in  most  specimens,  basal  foveae 
broad,  smooth  or  faintly  rugulose  or  punctulate, 
basal  angles  obtuse,  basal  sinuation  of  lateral 
margin  absent,  lateral  explanation  moderately 
wide  and  more  broadened  posteriorly  only,  mid- 
lateral  seta  absent;  elytral  silhouette  subovoid, 
slightly  narrowed  basally,  lateral  margins  slight- 
ly rounded;  hindwing  (Fig.  34)  reduced  in  length; 
hind  coxa  bi-  or  plurisetose  basally;  third  to  fifth 
visible  abdominal  sterna  each  with  two  or  more 
pairs  of  posterior  paramedial  setae;  specimen 
from  locality  in  Chuska  Mountains,  Arizona 
(Fig.  61). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Chuska  Mountains,  in 
which  the  type-locality  is  found. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  61; 
restricted  to  the  Chuska  Mountains,  Arizona.  I 
have  studied  specimens  from  the  following  lo- 
cality. 

United  States  of  America 

ARIZONA:  Apache  County,  Lukachukai  Creek  (at  Wagon 
Wheel  Campground  [2,260  m-2,320  m])  [July-Aug.]  (18;  CAS, 
MCZ,  UASM,  USNM). 

Nebria  darlingtoni,  new  species 

(Figures  20,  62) 
HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Calif.,  El 


Dorado  Co.,  Sierra  Nevada,  S.  F.  American  River.,  3  mi.  w. 
of  Riverton,  910m,  16July75  D.  H.  &  B.A.  Kavanaugh  Stop 
#15-29"!  "D.  H.  Kavanaugh  Collection"  [orange  label]/ 
"Holotype  Nebria  darlingtoni  Kavanaugh  det.  D.  H.  Kavan- 
augh 1976"  [red  label]/  "California  Academy  of  Sciences  Type 
No.  12499."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS. 
In  total,  170  PARATYPES  (105  males  and  65  females)  are  de- 
posited in  the  following  collections:  AMNH,  CAS,  DHKa, 
ICCM,  KSUC,  KUSM,  MCZ,  UASM,  USNM. 

TYPE-LOCALITY. — South  Fork  American  River  (3  miles  [ca. 
4.8  km]  w  of  Riverton),  El  Dorado  County,  California. 

DIAGNOSTIC  COMBINATION. — Body  black, 
legs  dark  brown  or  piceus;  microsculpture  on 
pronotum  and  elytra  very  deeply  impressed, 
meshes  isodiametric,  alveolae  convex;  prono- 
tum (Fig.  20)  with  lateral  explanation  narrow  at 
middle,  broadened  posteriorly,  basal  angles 
rectangular  or  slightly  acute,  basal  sinuation  of 
lateral  margin  moderately  short  and  deep,  mid- 
lateral  seta  absent;  elytra  with  humeral  carina 
slightly  developed,  slightly  projected  anteriorly, 
elytral  silhouette  ovoid,  markedly  narrowed  ba- 
sally, humeral  angles  moderately  rounded  and 
obtuse;  hind  coxa  bi-  or  plurisetose  basally; 
third  to  fifth  visible  abdominal  sterna  each  with 
two  or  more  pairs  of  posterior  paramedial  setae; 
specimen  from  Sierra  Nevada,  California  (Fig. 
62). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  species  in  honor  of  Philip 
J.  Darlington,  Jr.,  biogeographer,  student  of 
carabid  evolution,  and  friend,  whose  work  and 
personal  encouragement  have  provided  inspira- 
tion in  this  study. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  62; 
apparently  restricted  to  the  canyon  of  the  South 
Fork  of  the  American  River,  Sierra  Nevada, 
California.  I  have  studied  specimens  from  the 
following  localities. 

United  States  of  America 

CALIFORNIA:  El  Dorado  County  [Aug.]  (9;  CAS,  ICCM, 
KSUC,  MCZ,  USNM),  Kyburz  (2  miles  [ca.  3.2  km]  E  on 
South  Fork  American  River  [1,340  m])  [July]  (5;  CAS, 
DHKa),  Pacific  House  [Sep.]  (5;  CAS),  Riverton  (South  Fork 
American  River  [980  m],  3  miles  [ca.  4.8  km]  w  at  Bridal  Falls 
Picnic  Area  [910  m])  [July-Aug.]  (151;  AMNH,  CAS,  DHKa, 
KUSM,  MCZ,  UASM),  White  Hall  [June]  (2;  UASM). 

Nebria  navajo,  new  species 

(Figures  43,  63) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "Kayenta  Ariz.  19 
mi.  sw  VI-18-33"/  "Navajo  Co.  Alt.  6500  ft."/  "Ansel  F  Hall 
Exp.  1933"/  "Holotype  Nebria  navajo  Kavanaugh  det.  D.  H. 
Kavanaugh  1976"  [red  label]/  "California  Academy  of  Sci- 
ences Type  No.  12509."  One  male  PARATYPE  (same  data  as 
holotype)  also  in  CAS. 

TYPE-LOCALITY. — Kayenta  (19  miles  [ca.  31  km]  sw,  Na- 
vajo County,  Arizona. 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


105 


DIAGNOSTIC  COMBINATION. — Body  pale  tan- 
orange,  legs  pale  yellow;  microsculpture  on 
pronotum  and  elytra  moderately  impressed, 
meshes  isodiametric,  alveolae  flat;  pronotum 
with  lateral  explanation  narrow  at  middle, 
broadened  posteriorly,  basal  angles  rectangular 
or  slightly  acute,  basal  sinuation  of  lateral  mar- 
gin moderately  short  and  deep,  midlateral  seta 
absent;  elytra  with  humeral  carina  slightly  de- 
veloped, slightly  projected  anteriorly,  elytral  sil- 
houette ovoid,  markedly  narrowed  basally,  hu- 
meral angles  moderately  rounded  and  obtuse; 
hindwing  short,  narrow;  hind  coxa  bi-  or  plu- 
risetose  basally;  apex  of  fourth  tarsomere  of 
hind  tarsus  with  ventrolateral  lobe  long;  third  to 
fifth  visible  abdominal  sterna  each  with  two  or 
more  pairs  of  posterior  paramedial  setae;  male 
with  apex  of  median  lobe  as  in  Figure  43;  spec- 
imen from  Arizona  (Fig.  63). 

DERIVATION  OF  TAXON  NAME. — This  species 
is  named  for  the  people  native  to  the  region  in- 
cluding the  type-locality. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  63; 
known  only  from  the  type-locality  in  northeast- 
ern Arizona.  I  have  studied  specimens  from  the 
following  locality. 

United  States  of  America 

ARIZONA:  Navajo  County,  Kayenta  (19  miles  [ca.  31  km] 
sw  [1,980m])  [June]  (2:  CAS). 

Nebria  gebleri  cascadensis,  new  subspecies 

(Figures  21,  64) 

Nebria  gebleri;  auctoriim.  — HATCH  1939:118  (in  part). 
— LINDROTH  1961:83  (in  part).  —VAN  DYKE  1925:119  (in 
part);  1926: 10  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wash.,  Mt. 
Rainier  N.  P.,  Paradise  R.  above  Narada  Falls,  4580'-4800', 
8Aug72  DHKavanaugh  &HGoulet'V  "D.  H.  Kavanaugh  Col- 
lection" [orange  label]/  "Holotype  Nebria  gebleri  cascadensis 
Kavanaugh  det.  D.  H.  Kavanaugh  1976"  [red  label]/  "Cali- 
fornia Academy  of  Sciences  Type  No.  12502."  ALLOTYPE 
(same  data  as  holotype)  also  in  CAS.  In  total,  1696  PARATYPES 
(944  males  and  752  females)  are  deposited  in  the  following 
collections:  AMNH,  ANSP,  BFCa,  CAS,  CNC,  CUIC, 
DHKa,  DJLa,  FMNH,  INHS,  JSch,  KSUC,  KUSM,  LRus, 
MCZ,  MSU,  NMDo,  ODA,  OSUC,  OSUO,  PURC,  ROM, 
RTBe,  SDSU,  SJSC,  UAFA,  UASM,  UBC,  UCR,  UIMI, 
UMMZ,  USNM,  UWBM,  UWEM,  WSU. 

TYPE-LOCALITY. — Paradise  River  (above  Narada  Falls), 
Mount  Rainier  National  Park,  Washington. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size,  dark,  with  a  pair  of  pale  paramedial  spots 
on  vertex;  pronotum  (Fig.  21)  markedly  cordate, 
with  or  without  faint  violet  metallic  reflection, 
sinuation  of  lateral  margin  very  deep,  midlateral 


seta  present,  basolateral  seta  present;  elytra 
moderately  convex,  with  distinct  violet  or 
brassy  metallic  reflection,  elytral  silhouette  sub- 
ovoid,  slightly  narrowed  basally,  basal  margin- 
ation  moderately  concave,  humeral  angle  round- 
ly obtuse,  humeral  carina  moderately  to  markedly 
developed,  fifth  elytral  interval  without  setae  (in 
few  individuals  with  one  to  three  setae),  not  (or 
in  few  individuals  moderately)  catenate,  striae 
moderately  impressed;  specimen  from  locality 
in  Cascade  Range  of  central  Oregon  to  southern 
British  Columbia  (Fig.  64). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Cascade  Mountain  Range, 
in  which  members  of  this  taxon  are  widely  dis- 
tributed. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  64; 
restricted  to  the  Cascade  Range  (from  central 
Oregon  to  southern  British  Columbia),  the  ad- 
jacent mountain  ranges  west  of  the  Okanagan 
lowland  in  southwestern  British  Columbia,  and 
to  Vancouver  Island  (not  yet  recorded  from  the 
Olympic  Peninsula,  Washington,  although  I  pre- 
dict that  it  will  be  found  to  occur  there).  I  have 
studied  specimens  from  the  following  localities. 

Canada 

BRITISH  COLUMBIA:  Copper  Mountain  [Aug.]  (4:  UBC), 
Cultus  Lake  Provincial  Park  (Entrance  Bay  Campground) 
[June]  (4;  ROM),  Garibaldi  (Cheakamus  River)  [Sep.]  (2: 
JSpe),  Garibaldi  Provincial  Park  (Diamond  Head  Trail  [1,070 
m])  [July]  (3;  CNC),  Hope  [June-July]  (8;  BFCa,  IHNS), 
Hunter  Creek  (at  Restmore  Lodge)  [July]  (3:  CAS),  Manning 
Provincial  Park  (Blackwell  Peak  [910  m].  Lightning  Lake, 
Nicomen  Ridge,  Skagit  River)  [May-Sep.]  (35:  CAS,  CNC, 
DHKa,  KUSM,  MCZ,  MSU,  RTBe,  UBC,  UMMZ),  Midday 
Valley  (near  Merritt)  [Aug.]  (1;  INHS),  Nicolum  River  (at 
Hope  Trail)  [July]  (2;  UBC),  North  Bend  [July]  (19;  USNM), 
Skagit  River  (at  Hope  Trail  [760  m])  [July]  (3;  UBC),  Snass 
Creek  (at  Warburton  Trail  [760  m])  [July]  (5;  UBC),  Spious 
Creek  [May]  (1:  CAS),  Vancouver  area  (2:  ANSP),  Whipsaw 
Creek  (at  Hope  Trail  [1,220  m])  [May,  July]  (4:  UASM,  UBC); 
Vancouver  Island,  Port  Alberni  (Robertson  Creek  Spawning 
Channel)  [June]  (1;  ROM). 

United  States  of  America 

OREGON:  Clackamas  County,  Austin  Hot  Springs  [July]  (1; 
OSUO),  Brightwood  (Salmon  River)  [July]  (8;  DHKa),  Mount 
Hood  (s  slope  below  Timberline  Lodge  [1,190  m])  [Aug.]  (2; 
DHKa),  Rhododendron  ([490  m]  and  I  mile  [ca.  1.6  km]  SE  at 
Zig  Zag  River  [600  m])  [July-Aug.]  (55;  CAS,  DHKa,  SJSC, 
UWBM):  Deschutes  County,  Elk  Lake  [July]  (1:  UWBM). 
Upper  Deschutes  River  [May]  (1 :  UWBM);  Hood  River  Coun- 
ty, Hood  River  [May,  July]  (8;  CAS,  USNM),  Mount  Hood 
(Cloud  Cap,  Hood  River  Meadows  Ski  Area  [1,710  m].  Sand 
Creek,  Tillie  Jane  Creek,  Umbrella  Falls)  [June-Sep.[  (70; 
CAS,  DHKa,  MCZ,  NMDo,  ODA,  UCR,  UIMI,  UWBM): 
Marion  County,  Detroit  ([460  m]  and  5  miles  [ca.  8.0  km]  N) 
[July-Aug.]  (3;  LRus,  UWBM);  Multnomah  County,  Horsetail 
Falls  ([120  m])  [May-July]  (87;  DHKa,  JSch,  ODA,  OSUO). 
WASHINGTON:  (32;  ANSP,  CAS,  INHS,  MCZ,  OSUC, 


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USNM);  Chelan  County,  Fish  Lake  [July]  (1;  UWBM),  Leav- 
enworth  [July]  (8;  ANSP,  KUSM,  MCZ,  SDSU,  USNM), 
Tumwater  Canyon  ([610  m])  [May]  (1;  LRus);  Cowlitz 
County,  Ariel  [May]  (1;  UWBM],  Cougar  [Sep.]  (2;  OSUO), 
Hoffstadt  Creek  (at  Toutle  River)  [July]  (2;  MSU,  OSUO), 
North  Fork  Toutle  River  (14  miles  [ca.  23  km]  E  of  Toutle 
[300  m])  [Aug.]  (33;  CAS,  DHKa),  Toutle  River  [July]  (11; 
UWBM);  King  County,  Baring  [July]  (2;  FMNH),  Green 
River  Gorge  (Green  River)  [May,  Aug.]  (5;  UWBM),  Green- 
water  River  (at  Greenwater  Campground)  [June]  (1;  UWBM), 
North  Bend  [July]  (17;  CAS,  CUIC,  MCZ),  Renton  (Cedar 
River)  [May]  (1;  UWBM),  Skykomish  [July,  Sep.]  (2;  UASM, 
UWBM),  Snoqualmie  [June]  (4;  UIMI),  Stevens  Pass  (and  4 
miles  [ca.  6.4  km]  w)  [June,  Aug.]  (5;  LRus,  UASM),  Welling- 
ton [July]  (10;  CAS,  MCZ,  USNM,  UWEM),  White  River  (5 
miles  [ca.  8  km]  w  of  Greenwater  [1,490  m],  Slippery  Creek) 
[July-Aug.]  (42;  DHKa,  UWBM);  Kittitas  County,  Blewett 
Pass  (Niger  Creek)  [July]  (2;  USNM),  Easton  (22;  CAS, 
INHS,  MCZ,  USNM),  Snoqualmie  Pass  [July]  (4;  CAS), 
Teanaway  Ridge  ([910  m])  [July]  (10;  LRus);  Klickitat  County, 
Klickitat  Valley  [July]  (1;  MCZ);  Lewis  County,  Lake  Creek 
(3  miles  [ca.  4.8  km]  NE  of  Packwood  [370  m])  [July]  (5; 
DHKa),  White  Pass  (and  1.5  miles  [ca.  2.4  km]  w  at  Millridge 
Creek  [1,280  m])  [June-July]  (42;  DHKa,  SJSC,  USNM); 
Mount  Rainier  National  Park  ([760  m-1,830  m])  [June-Sep.] 
(30;  ANSP,  CAS,  CUIC,  KSUC,  MCZ,  OSUO,  SJSC,  UIMI, 
UWBM),  Bear  Flat  ([1,520  m-1,830  m])  [July]  (1;  CAS), 
Carbon  River  [June]  (4;  UWBM),  Cayuse  Pass  [1,400  m] 
[June]  (4;  USNM),  Fish  Creek  (at  West  End  Road)  [June] 
(1;  ROM),  Fryingpan  Creek  ([1,160  m-1,190  m])  [July-Aug.] 
(38;  CNC,  DHKa,  SJSC),  Horse  Creek  (near  Long- 
mire)  [July]  (15;  CAS),  Ipsut  Creek  Campground 
[Aug.]  (1;  OSUO),  Kautz  Creek  (at  Nisqually  Riv- 
er) [Sep.]  (2;  UWBM),  Longmire  ([760  m])  [June-Aug.] 
(25;  CAS,  SJSC,  USNM),  Narada  Falls  ([1,370  m-1,460  m]) 
[July-Aug.]  (118;  DHKa,  USNM,  UWBM),  Nisqually  River 
(0.1  miles  [ca.  0.2  km]  above  Paradise  River  [980  m],  toe  of 
Nisqually  Glacier  [1,220  m-1,520  m])  [July-Aug.]  (7;  AMNH, 
CAS,  CUIC,  DHKa,  MCZ),  North  Fork  Puyallup  River  (be- 
low toe  of  Puyallup  Glacier  [1,100  m-1,650  m])  [July-Aug.] 
(22;  CNC,  DHKa,  USNM),  Ohanapecosh  [June,  Aug.]  (18; 
DJLa,  LRus,  SJSC,  UIMI,  USNM),  Paradise  area  (below  Slu- 
skin  Falls,  Paradise  Park  [1,220  m-1,830  m],  Paradise  River 
[1,400  m-1,680  m].  Paradise  Valley)  [June-Sep.]  (88;  CAS, 
CUIC,  DHKa,  FMNH,  MCZ,  USNM,  UWBM),  St.  Andrews 
Creek  (at  West  End  Road  [1,160  m])  [Aug.]  (5;  DHKa),  Silver 
Creek  [Aug.]  (10;  UWBM),  Sluskin  Falls  [July]  (8;  UWBM), 
Stevens  Creek  ([1,190  m-1,220  m])  [June-July]  (4;  DHKa, 
USNM),  Sunbeam  Falls  ([1,220  m])  [May]  (1;  CNC),  Sunrise 
area  [July]  (2;  LRus),  Tahoma  Creek  ([910  m])  [July]  (1; 
USNM),  West  End  Road  [July]  (1;  CAS),  White  River  ([1,190 
m]  and  at  White  River  Campground  [1,520  m])  [July-Aug.] 
(12;  FMNH,  SJSC,  USNM,  UWBM),  Van  Trump  Creek 
(above  Christine  Falls  [1,220  m- 1,280  m])  [July-Aug.]  (62; 
DHKa,  UWBM);  Pierce  County.  Goat  Creek  (at  Nisqually 
River  [610  m])  [July]  (130;  DHKa),  Fairfax  [Oct.]  (1;  FMNH), 
Poch  Creek  (at  Carbon  River)  [Aug.]  (5;  UCB,  UWBM),  Ten- 
as  Creek  (5  miles  [ca.  8.0  km]  E  of  Ashford  [610  m])  [July]  (1; 
DHKa),  White  River  (at  Silver  Springs  Campground  [790  m]) 
[June,  Aug.]  (15;  DHKa,  USNM);  Skamania  County,  Carson 
[Aug.]  (6;  UIMI),  Council  Lake  (SE  on  Forest  Road  123  [1,260 
m])  [Aug.]  (4:  UASM),  Mount  St.  Helens  (Maratta  Creek) 
[July]  (3;  DHKa),  North  Fork  Cipsus  River  [June]  (14; 
UWBM),  North  Fork  Toutle  River  (at  Spirit  Lake  Lodge  [940 


m])  [Aug.]  (31;  CAS,  DHKa),  Spirit  Lake  [July,  Sep.]  (3; 
DHKa,  UWBM),  Stabler  [June]  (1;  OSUO),  Wind  River  (8 
miles  [ca.  13  km]  N  of  Carson  [340  m])  [May,  July]  (14; 
DHKa);  Snohomish  County,  Darrington  (Clear  Creek  Falls) 
[May]  (5;  UWBM).  Glacier  Peak  [July-Aug.]  (2;  CAS, 
UWBM);  Whatcom  County,  Glacier  (North  Fork  Nooksack 
River)  [July]  (13:  CAS,  LRus),  Lookout  Creek  (4  miles  [ca. 
6.4  km]  E  of  Glacier  [380  m])  [Aug.]  (18;  CAS,  DHKa),  Mount 
Baker  (Austin  Pass  [1,520  m],  Bagley  Creek  [670  m],  Razor 
Hone  Creek  [1,680  m])  [July-Sep.]  (125:  CAS,  DHKa,  JSpe, 
LRus,  UCR,  UWBM),  North  Fork  Nooksack  River  (4  miles 
[ca.  6.4  km]  E  of  Glacier  [380  m],  Silver  Fir  Campground  [610 
m])  [Aug.]  (3;  CAS,  DHKa),  Shuksan  ([760  m])  [Aug.]  (3; 
LRus),  Shuksan  Creek  [Aug.]  (12;  UWBM),  Skagit  River  (at 
Newhalem)  [July]  (2;  UWBM);  Yakima  County,  American 
River  (at  Hell's  Crossing  [1,040  m])  [July-Aug.]  (17;  DHKa, 
ICCM,  KUSM,  MCZ,  UIMI,  UMMZ,  UWBM),  Bumping 
River  (0.8  miles  [ca.  1.3  km]  sw  of  Highway  410  [1,010  m]) 
[July-Sep.]  (92;  CAS,  DHKa,  ICCM,  MCZ,  MSU,  PURC, 
UAFA,  USNM,  WSU),  Dog  Lake  ([1,310  m])  [July]  (25; 
DHKa),  Mount  Adams  (Bird  Creek  [910  m-2,440  m].  Chap- 
arral Creek  [880  m],  Klickitat  River)  [June-July,  Sep.]  (82; 
AMNH,  ANSP,  CAS,  CUIC,  FMNH,  ICCM,  MCZ,  UASM, 
UIMI,  USNM,  UWBM,  WSU),  Naches  River  [July]  (2; 
USNM),  Tampico  [July]  (1;  USNM),  Tieton  Dam  [Aug.]  (2; 
AMNH,  WSU),  Wildcat  Creek  (at  Tieton  River  [880  m])  [July] 
(3;  DHKa,  OSUO),  Yakima  area  (11:  JSch,  MCZ,  USNM, 
WSU),  Yakima  Indian  Reservation  ([910  m])  [July]  (1;  CAS). 

Doubtful  Records 

UNITED  STATES  OF  AMERICA — COLORADO:  Leavenworth 
Valley  (1;  ANSP).  WASHINGTON:  Pullman  (1;  ICCM). 

Specimens  Without  Locality  Data:  (4:  CDA,  MCZ,  MSU, 
UWBM). 

Nebria  gebleri  fragariae,  new  subspecies 

(Figures  22,  44,  49,  64) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Ore.,  Grant 
Co.,  Blue  Mts.,  Strawberry  Cr.,  5800'  28  May  1971,  DHKa- 
vanaugh  &  EAMartinko'V  "D.  H.  Kavanaugh  Collection" 
[orange  label]/  "Holotype  Nebria  gebleri  fragariae  Kavanaugh 
det.  D.  H.  Kavanaugh  1979"  [red  label]/  "California  Academy 
of  Sciences  Type  No.  12501."  ALLOTYPE  (same  data  as  ho- 
lotype)  also  in  CAS.  In  total,  27  PARATYPES  (21  males  and  6 
females)  are  deposited  in  the  following  collections:  CAS, 
DHKa,  UASM,  USNM. 

TYPE-LOCALITY. — Strawberry  Creek  (1,770  m).  Grant 
County,  Oregon. 

DIAGNOSTIC  COMBINATION. — Size  moderate, 
standardized  body  length  male  less  than  10.5 
mm,  female  less  than  11.0  mm;  head  moderate 
in  size,  dark,  with  a  pair  of  pale  paramedial  spots 
on  vertex;  pronotum  (Fig.  22)  moderately  cor- 
date, with  or  without  faint  green  metallic  reflec- 
tion, midlateral  seta  present,  basolateral  seta 
present;  elytra  with  distinct  green  metallic  re- 
flection, elytral  silhouette  subrectangular,  broad 
basally;  male  with  apex  of  median  lobe  as  in 
Figure  44;  female  with  anterodorsal  sclerite  in 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


107 


bursa  copulatrix  (Fig.  49);  specimen  from  local- 
ity in  Strawberry  Range,  Blue  Mountains,  east- 
ern Oregon  (Fig.  64). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Strawberry  Range  of  the 
Blue  Mountains,  in  which  the  type-locality  is 
found.  The  subspecific  epithet  is  a  form  of  the 
scientific  name  for  the  strawberry  genus,  Fra- 
garia  Linnaeus. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  64; 
restricted  to  the  Strawberry  Range  of  the  Blue 
Mountains  of  eastern  Oregon.  I  have  studied 
specimens  from  the  following  locality. 

United  States  of  America 

OREGON:  Grant  County,  Strawberry  Creek  ([1,770  m]) 
[May]  (29:  CAS,  DHKa,  UASM,  USNM). 

Nebria  gebleri  siskiyouensis,  new  subspecies 

(Figures  23,  64) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  •'U.S.A.,  Calif.,  Trin- 
ity Co.,  Trinity  Alps,  S.  F.  Salmon  R.  at  Big  Flat  Cmpgd., 
1490m,  18July75  D.  H.  &  B.  A.  Kavanaugh  Stop  #75-36' V 
"D.  H.  Kavanaugh  Collection"  [orange  label]/  "Holotype 
Nebria  gebleri  siskiyouensis  Kavanaugh  det.  D.  H.  Kavan- 
augh 1976"  [red  label]/ "California  Academy  of  Sciences  Type 
No.  12503."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS. 
In  total,  33  PARATYPES  (19  males  and  14  females)  are  depos- 
ited in  the  following  collections:  CAS,  DHKa,  FMNH, 
UASM.  USNM. 

TYPE-LOCALITY. — South  Fork  Salmon  River  (at  Big  Flat 
Campground),  Trinity  County,  California. 

DIAGNOSTIC  COMBINATION. — Head  dark,  with 
a  pair  of  pale  paramedial  spots  on  vertex;  prono- 
tum  (Fig.  23)  with  apical  angles  moderately  nar- 
row, midlateral  seta  present,  basolateral  seta 
present;  elytra  without  metallic  reflection,  ely- 
tral  silhouette  subovoid,  relatively  short  and 
broad;  hindwing  full-sized;  specimen  from  lo- 
cality in  Klamath  Mountains  system  of  Coast 
Ranges  in  northwestern  California  or  south- 
western Oregon  (Fig.  64). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Siskiyou  Mountains,  in 
which  members  of  this  taxon  were  first  collect- 
ed. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  64; 
restricted  to  the  Klamath  Mountains  system  of 
the  Coast  Ranges  in  northwestern  California  and 
southwestern  Oregon.  I  have  studied  specimens 
from  the  following  localities. 

United  States  of  America 

CALIFORNIA:  Trinity  County,  Boulder  Creek  (at  Goldfield 
Campground  [1,070  m])  [July]  (2:  CAS,  DHKa),  South  Fork 


Salmon  River  (at  Big  Flat  Campground  [1,490  m])  [June-July] 
(30;  CAS,  DHKa,  FMNH,  UASM,  USNM),  Taylor  Lake 
Road  (branch  of  Taylor  Creek  [1,750  m])  [Aug.]  (1;  CAS). 
OREGON:  Jackson  County,  Little  Applegate  River  ([700  m]) 
[Aug.]  (2;  CAS,  FMNH). 

Nebria  carri,  new  species 

(Figures  10,  24,  65) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Ida.,  Blaine 
Co.,  Sawtooth  Mts.,  Smoky  Range,  2  mi.  e.  Dollarhide  Sum- 
mit, N.  F.  Warm  Springs  Creek,  7700'-7900',  21  Aug. 73 
DHKavanaugh  Family"/  "D.  H.  Kavanaugh  Collection"  [or- 
ange label]/  "Holotype  Nebria  carri  Kavanaugh  det.  D.  H. 
Kavanaugh  1976"  [red  label]/  "California  Academy  of  Sci- 
ences Type  No.  12498."  ALLOTYPE  (same  data  as  holotype) 
also  in  CAS.  In  total,  152  PARATYPES  (63  males  and  89  fe- 
males) are  deposited  in  the  following  collections:  BFCa,  CAS, 
DHKa,  FMNH,  GASh,  MCZ,  UASM,  USNM. 

TYPE-LOCALITY. — Dollarhide  Summit,  Blaine  County,  Ida- 
ho. 

DIAGNOSTIC  COMBINATION. — Head  dark,  with 
a  pair  of  pale  paramedial  spots  on  vertex;  pen- 
ultimate labial  palpomere  bisetose;  mentum 
(Fig.  10)  with  M:!  setae  absent;  pronotum  (Fig. 
24)  with  midlateral  seta  present,  basolateral  seta 
absent;  specimen  from  locality  east  of  Columbia 
Basin  (Fig.  65). 

DERIVATION  OF  TAXON  NAME. — I  take  plea- 
sure in  naming  this  species  in  honor  of  John  and 
Betty  Carr  (Calgary,  Alberta),  who  first  collect- 
ed specimens  of  same,  and  who  have  provided 
help  and  encouragement  to  me  and  to  other  stu- 
dents during  our  respective  studies. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  65; 
known  at  present  only  from  ranges  of  the  Saw- 
tooth Mountains  system  and  from  the  Seven 
Devils  Mountains  of  southcentral  and  western 
Idaho,  respectively.  I  have  studied  specimens 
from  the  following  localities. 

United  States  of  America 

IDAHO:  Adams  County,  Towsley  Springs  (9  miles  [ca.  14 
km]  NNE  of  Cuprum  [2,070  m])  [July-Aug.]  (4;  BFCa,  DHKa): 
Blaine  County,  Dollarhide  Summit  (1.0  to  2.0  miles  [ca.  1.6 
to  3.2  km]  E  on  North  Fork  Warm  Springs  Creek  [2,350  m- 
2,500  m])  [Aug.]  (138:  CAS,  DHKa,  MCZ.  UASM,  USNM), 
Twin  Lakes  Trail  (w  of  Petit  Lake)  [July]  (2:  BFCa);  Camas 
County,  Carrie  Creek  (32  miles  [ca.  51  km]  wsw  of  Ketchum 
[2,400  m])  [Aug.]  (9;  DHKa):  Elmore  County,  Trinity  Lakes 
[Aug.]  (1:  GASh). 

Nebria  kincaidi  balli,  new  subspecies 

(Figures  11,  31,65) 

Nebria  kincaidi;  auctorum.  — BLACKWELDER  AND  BLACK- 
WELDER  1948:2  (in  part).  — ERWIN  AND  BALL  1972:85  (in 
part).  —HATCH  1939:121  (in  part).  — LINDROTH  1961:88 
(in  part).  —VAN  DYKE  1919:8  (in  part). 


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Nebria   coltimbiana;   auctorum   (not   Casey).   — HATCH 
1939:121  (in  part).  —VAN  DYKE  1919:8  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wash.,  Mt. 
Rainier  N.  P.,  Paradise  R.  above  Narada  Falls,  4580'-4800', 
8Aug72  DHKavanaugh  &  HGouIefV  "D.  H.  Kavanaugh  Col- 
lection" [orange  label]/  "Holotype  Nebria  kincaidi  balli  Ka- 
vanaugh del.  D.  H.  Kavanaugh  1976"  [red  label]/  "California 
Academy  of  Sciences  Type  No.  12505."  ALLOTYPE  (same 
data  as  holotype)  also  in  CAS.  In  total,  366  PARATYPES  (18$. 
males  and  177  females)  are  deposited  in  the  following  collec- 
tions: AMNH,  CAS,  CNC,  CUIC,  DHKa,  FMMH,  KUSM, 
LACM,  LRus,  MCZ,  MSU,  ODA,  OSUO,  SJSC,  UASM, 
USNM,  UWBM,  WSU. 

TYPE-LOCALITY. — Paradise  River  (above  Narada  Falls), 
Mount  Rainier  National  Park,  Washington. 

DIAGNOSTIC  COMBINATION. — Head  dark,  with 
a  pair  of  pale  paramedial  spots  on  vertex;  pen- 
ultimate labial  palpomere  trisetose;  mentum 
(Fig.  11)  with  M:,  setae  present;  pronotum  with 
midlateral  seta  present,  basolateral  seta  absent; 
elytra  with  basal  margination  (Fig.  31)  moder- 
ately long,  smooth  or  faintly  wavy,  slightly  to 
moderately  concave;  specimen  from  Cascade 
Range  of  central  Washington  to  northern  Oregon 
(Fig.  65). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  subspecies  in  honor  of 
George  E.  Ball,  my  mentor  and  friend. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  65; 
restricted  to  the  Cascade  Range  of  central 
Washington  and  northern  Oregon.  Present 
known  range  extends  from  Mount  Rainier 
(Washington)  south  to  Mount  Hood  (Oregon). 
I  have  studied  specimens  from  the  following  lo- 
calities. 

United  States  of  America 

OREGON:  Hood  River  County,  Mount  Hood  ([910-1,830  m], 
Cloud  Cap  Road  at  Tillie  Jane  Road,  Hood  River  Meadows 
Ski  Area  [1,620  m-1,710  m],  Tillie  Jane  Creek  [1,830  m],  Um- 
brella Falls)  [June-Sep.]  (67:  AMNH,  CAS,  CUIC,  DHKa, 
MCZ,  MSU,  ODA,  USNM,  UWBM,  WSU).  WASHINGTON: 
Mount  Ranier  National  Park  [July-Sep.]  (4;  CUIC,  SJSC, 
WSU),  Cayuse  Pass  ([1,400  m])  [June]  (1;  USNM),  Fryingpan 
Creek  ([1,160  m-1,190  m])  [July-Aug.]  (2;  CNC,  DHKa),  In- 
dian Henry's  ([1,370  m-1,520  m])  [July]  (1:  UASM),  Louise 
Lake  [July]  (5;  UASM),  Mowich  Lake  [July]  (4;  OSUO, 
UWBM),  Mystic  Lake  [July]  (1;  WSU),  Nisqually  River 
([1,220  m])  [May]  (4;  CNC),  Paradise  area  (above  Narada  Falls 
[1,520  m],  Paradise  Park  [1,520  m- 1,830  m],  Paradise  River 
[1,460  m-1,830  m])  [June-Aug.]  (265;  CAS,  CNC,  CUIC, 
DHKa,  FMNH,  KUSM,  LRus,  MCZ,  OSUO,  UASM, 
USNM,  UWBM),  Ricksecker  Point  [Sep.]  (1;  UWBM),  Slu- 
skin  Falls  [July]  (1:  UWBM),  Snow  Lake  ([1,520  m])  [Aug.] 
(1;  CNC),  Spray  Park  ([1,980  m])  [Aug.]  (1:  LRus),  Sunrise 
area  [July]  (2;  CAS,  LRus),  Van  Trump  Creek  (above  Chris- 
tine Falls  [1,220  m-1,280  m])  [July]  (7;  DHKa),  Van  Trump 
Park  ([1,830  m])  [Aug.]  (1;  CNC),  West  End  Drive  [July]  (1; 
CAS);  Pierce  County,  Coplay  Lake  [May]  (1;  UWBM);  Ska- 


mania  County,   Spirit   Lake  ([910  m-1,770  m])   [Aug.]  (1; 
OSUO). 

Doubtful  Records 

UNITED  STATES  OF  AMERICA — OREGON:  Deschutes  Coun- 
ty, Bend  [July]  (8:  LACM). 

Nebria  spatulata  sierrae,  new  subspecies 

(Figures  6,  66) 

Nebria  spatulata;  ERWIN  AND  BALL  1972:83  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Calif., 
Mono  Co.,  Sierra  Nevada  Mts.,  ne.  face  White  Mt.,  along 
stream  in  cirque  above  Big  Horn  Lake,  3290m-3480m,  21  July 
74  D.H. Kavanaugh  Collector"/  "D.  H.  Kavanaugh  Collec- 
tion" [orange  label]/  "Holotype  Nebria  spatulata  sierrae  Ka- 
vanaugh det.  D.  H.  Kavanaugh  1976"  [red  label]/  "California 
Academy  of  Sciences  Type  No.  12516."  ALLOTYPE  (same 
data  as  holotype)  also  in  CAS.  In  total,  58  PARATYPES  (28 
males  and  30  females)  are  deposited  in  the  following  collec- 
tions: CAS,  DHKa,  MCZ,  RPPa,  UASM,  UCB,  USNM. 

TYPE-LOCALITY.— White  Mountain  (east  slope,  above  Big 
Horn  Lake),  Mono  County,  California. 

DIAGNOSTIC  COMBINATION. — Size  moderate: 
standardized  body  length  male  less  than  12.0 
mm,  female  less  than  12.4  mm;  head  with  a  pair 
of  pale  paramedial  spots  (or  pair  fused  to  form 
a  single  median  spot),  genae  and  occiput  not 
swollen,  eyes  moderate  in  size  and  convexity; 
antennal  scape  (Fig.  6)  moderate  in  length, 
slightly  arcuate,  markedly  narrowed  basally; 
pronotum  with  midlateral  seta  present,  basolat- 
eral seta  present;  elytra  rufous  or  piceus,  very 
dull,  with  microsculpture  deeply  impressed, 
meshes  isodiametric,  without  metallic  reflec- 
tion, elytral  silhouette  markedly  ovoid,  nar- 
rowed basally;  hindwing  vestigial;  specimen 
from  locality  on  Eastern  Divide  or  more  north- 
ern portion  of  Sierra  Nevada  (Fig.  66). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  Sierra  Nevada,  California, 
to  which  its  geographical  range  is  restricted. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  66; 
restricted  to  the  Eastern  Divide  of  the  Sierra 
Nevada  in  California;  presently  known  from  as 
far  north  as  Sonora  Pass  (Tuolumne  County)  and 
as  far  south  as  Big  Pine  Creek  (Inyo  County). 
I  have  studied  specimens  from  the  following  lo- 
calities. 

United  States  of  America 

CALIFORNIA:  Inyo  County,  Big  Pine  Creek  (at  Finger  Lake 
[3,290  m])  [Aug.]  (4;  CAS);  Mono  County,  H.  M.  Hall  Natural 
Area  ([3,350  m])  [July-Sep.]  (9;  RPPa),  Saddlebag  Lake  [Sep.] 
(3;  CAS),  White  Mountain  (NE  slope  in  cirque  above  Big  Horn 
Lake  [3,290  m-3,480  m])  [July]  (13;  DHKa);  Tuolumne  Coun- 
ty, Sonora  Pass  ([2,740  m-3,350  m])  [July]  (3:  UCB,  USNM); 
Yosemite  National  Park,  Maclure  Creek  Basin  ([3,290  m- 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


109 


3,350  m])  [Sep.]  (1;  DHKa),  Mount  Lyell  ([3,350  m|)  [July- 
Aug.]  (23;  CAS,  MCZ,  UASM),  Tioga  Pass  ([3,050  m[)  [Aug.] 
(2;  CAS). 

Doubtful  Records 

UNITED  STATES  OF  AMERICA — CALIFORNIA:  San  Bernar- 
dino County  [July]  (2:  CAS). 

Nebria  meanyi  lamarckensis,  new  subspecies 

(Figures  25,  32,  67) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Calif.,  Inyo 
Co.,  Sierra  Nevada,  Lamarck  Cr.  above  Upper  Lamarck  Lk., 
10700'-!  1000'  6July71  DHKavanaugh  &  EAMartinko"/  "D. 
H.  Kavanaugh  Collection"  [orange  label]/  "Holotype  Nebria 
meanyi  lamarckensis  Kavanaugh  del.  D.  H.  Kavanaugh  1976" 
[red  label]/  "California  Academy  of  Sciences  Type  No. 
12507."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS.  In 
total,  21  PARATYPES  (6  males  and  15  females)  are  deposited 
in  the  following  collections:  CAS.  DHKa,  UASM. 

TYPE-LOCALITY. —  Lamarck  Creek  (above  Upper  Lamarck 
Lake),  Inyo  County,  California. 

DIAGNOSTIC  COMBINATION. — Size  moderate, 
standardized  body  length  male  less  than  12.0 
mm,  female  9.5  to  12.3  mm;  head  dark,  with  a 
pair  of  pale  paramedial  spots  on  vertex;  prono- 
tum  (Fig.  25)  markedly  cordate,  basal  angles 
acute,  projected  posteriorly,  midlateral  seta 
present,  basolateral  seta  present;  elytra  with 
faint  or  moderate  violet,  blue,  or  green  metallic 
reflection,  elytral  silhouette  subrectangular, 
broad  basally,  basal  margination  (Fig.  32) 
straight  or  slightly  concave;  specimen  from  lo- 
cality in  southern  Sierra  Nevada  (California) 
(Fig.  67). 

DERIVATION  OF  TAXON  NAME. — The  name 
for  this  subspecies  refers  to  the  type-locality, 
Lamarck  Creek,  which  in  turn  refers  to  Jean 
Baptiste  Pierre  Antoine  Lamarck,  the  noted 
French  naturalist  and  pioneer  evolutionist. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  67; 
restricted  to  the  east  slope  of  the  southern  Sierra 
Nevada.  I  have  studied  specimens  from  the  fol- 
lowing localities. 

United  States  of  America 

CALIFORNIA:  Inyo  County,  Big  Pine  Creek  (below  Finger 
Lake  [3,140  m])  [Aug.]  (2:  CAS),  Lamarck  Creek  (above  Up- 
per Lamarck  Lake  [3,260  m-3,350  m])  [July]  (21:  CAS, 
DHKa,  UASM). 

Nebria  meanyi  sylvatica,  new  subspecies 

(Figures  26,  67) 

Nebria  meanyi;  HATCH  1939:118  (in  part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Wash., 
Olympic  N.  P.,  Olympic  Mts.,  Boulder  Cr.  at  Olympic  Hot 
Springs,  610m,  15Aug.74  D.  H.  Kavanaugh"/  "D.  H.  Kavan- 
augh Collection"  [orange  label]/  "Holotype  Nebria  meanyi 


sylvatica  Kavanaugh  del.  D.  H.  Kavanaugh  1976"  [red  label)/ 
"California  Academy  of  Sciences  Type  No.  12508."  ALLO- 
TYPE (same  data  as  holotype)  also  in  CAS.  In  total,  177  PARA- 
TYPES  (83  males  and  94  females)  are  deposited  in  the  following 
collections:  CAS,  CNC,  CUIC,  DHKa,  ICCM,  JSch.  UASM, 
USNM.  UWBM. 

TYPE-LOCALITY. — Boulder  Creek  (at  Olympic  Hot  Springs), 
Olympic  National  Park,  Washington. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  width,  dark,  with  a  pair  of  pale  par- 
amedial spots  on  vertex;  pronotum  (Fig.  26) 
markedly  cordate,  basal  sinuation  of  lateral  mar- 
gin very  short  and  deep,  midlateral  seta  present, 
basolateral  seta  present;  elytra  moderately  con- 
vex, elytral  silhouette  subrectangular  to  sub- 
ovoid,  moderately  broad  basally,  basal  margin- 
ation slightly  short,  humeral  angles  faintly  or 
moderately  obtuse,  humeral  carina  present, 
slightly  to  markedly  developed,  elytral  intervals 
markedly  convex,  fifth  interval  with  three  to  six 
setae,  markedly  catenate,  striae  deeply  im- 
pressed; specimen  from  locality  on  Vancouver 
Island  (British  Columbia)  or  Olympic  Peninsula 
(Washington)  (Fig.  67). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies name  is  derived  from  the  Latin  word,  syl- 
vaticus,  meaning  "of  the  forest" — a  reference 
to  the  deep  forest  habitat  occupied  by  taxon 
members. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  67; 
restricted  to  Vancouver  Island,  British  Colum- 
bia, and  the  Olympic  Peninsula,  Washington.  I 
have  studied  specimens  from  the  following  lo- 
calities. 

Canada 

BRITISH  COLUMBIA:  Vancouver  Island,  Alberni  (38  miles 
[ca.  61  km]  w)  [May](l;  CNC). 

United  States  of  America 

WASHINGTON:  Olympic  National  Park  [Aug.]  (11;  ICCM, 
MCZ,  UASM,  USNM),  Elwha  River  (at  Elwha  Campground) 
[Aug.]  (1;  CUIC),  Dosewallips  River  (at  Muscott  Camp- 
ground) [July]  (3;  UWBM),  Olympic  Hot  Springs  (Boulder 
Creek  [670  m-760  m])  [May-Aug.]  (160;  CAS,  CNC,  DHKa, 
JSch,  UWBM),  Sol  Due  Hot  Springs  [July]  (3:  JSch,  UWBM). 

Nebria  vandykei  wyeast,  new  subspecies 

(Figures  7.  68) 

Nebria  vandykei;  auctorum.  — ERWIN  AND  BALL  1972:87  (in 
part). —HATCH  1939:118  (in  part).  — LINDROTH  1961:86  (in 
part). 

Nebria  trifaria  vandykei  BANNINGER.  — HATCH  1953:56  (in 
part). 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A..  Ore.,  Clack- 
amas  Co.,  Cascade  Range,  s.  slope  Mt.  Hood,  headwaters 
Salmon  R.  near  Timberline  Lodge,  1830m-1950m,  3  Aug. 74 


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D.  H.  Kavanaugh"/  "D.  H.  Kavanaugh  Collection"  [orange 
label]/  "Holotype  Nebria  vandykei  wyeast  Kavanaugh  det.  D. 
H.  Kavanaugh  1976"  [red  label]/  "California  Academy  of  Sci- 
ences Type  No.  12517."  ALLOTYPE  (same  data  as  holotype) 
also  in  CAS.  In  total,  186  PARATYPES  (104  males  and  82  fe- 
males) are  deposited  in  the  following  collections:  CAS, 
DHKa,  MCZ,  OSUO,  UASM,  USNM.  UWBM. 

TYPE-LOCALITY. — Salmon  River  headwaters.  Mount  Hood, 
Clackamas  County,  Oregon. 

DIAGNOSTIC  COMBINATION. — Size  very  large, 
standardized  body  length  male  greater  than  12.0 
mm,  female  greater  than  12.5  mm;  head  dark, 
with  a  pair  of  pale  paramedial  spots  on  vertex; 
antennal  scape  (Fig.  7)  narrowed  basally, 
straight  or  slightly  arcuate;  pronotum  with  mid- 
lateral  seta  present,  basolateral  seta  present; 
elytra  without  metallic  reflection,  elytral  silhou- 
ette markedly  ovoid;  specimen  from  locality  in 
Cascade  Range  south  of  Columbia  River  (Fig. 
68). 

DERIVATION  OF  TAXON  NAME. — For  this  sub- 
species, I  have  selected  that  name  given  to 
Mount  Hood  by  local  native  peoples — Wy'east, 
which  means  "The  Mountain." 

GEOGRAPHICAL  DISTRIBUTION. — Figure  68; 
restricted  to  the  Cascade  Range  south  of  the 
Columbia  River;  present  known  range  from 
Mount  Hood  south  to  the  Three  Sisters  area 
(Oregon).  I  have  studied  specimens  from  the  fol- 
lowing localities. 

United  States  of  America 

OREGON:  Deschutes  County,  Middle  Sister  Peak  (E  slope  at 
North  Fork  Squaw  Creek  [l,950m-2,190  m])  [Aug.]  (19;  CAS, 
DHKa);  Hood  River  County,  Mount  Hood  ([1,740  m],  Hood 
River  Meadows  Ski  Area  [1,920  m-1,950  m],  headwaters  of 
Salmon  River  near  Timberline  Lodge  [1,830  m-1,950  m].  Sand 
Creek)  [June-Aug.]  (130:  CAS,  DHKa,  UWBM);  Marion 
County,  Mount  Jefferson  (N  slope  [1,860  m-1,920  m],  s  end 
of  Jefferson  Park  [1,830  m],  Russell  Creek  [1,680  m- 1,740  m], 
West  Branch  Whitewater  River  [1,830  m-1,890  m])  [Aug.]  (39; 
CAS,  DHKa,  OSUO). 

Nebria  schwarzi  beverlianna,  new  subspecies 

(Figures  1,  27,  36,  69) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.,  Wyo.,  Sub- 
lette  Co.,  Gros  Ventre  Mts.,  Hwy.  187/189,  8  mi.  nw.  Bon- 
durant,  Hoback  R.,  6900',  1-2  August  73  DHKavanaugh  Fam- 
ily"/ "D.  H.  Kavanaugh  Collection"  [orange  label]/  "Holotype 
Nebria  schwarzi  beverlianna  Kavanaugh  det.  D.  H.  Kavan- 
augh 1976"  [red  label]/ "California  Academy  of  Sciences  Type 
No.  12515."  ALLOTYPE  (same  data  as  holotype)  also  in  CAS. 
In  total,  254  PARATYPES  (133  males  and  121  females)  are  de- 
posited in  the  following  collections:  CAS,  DHKa,  FMNH, 
MCZ,  ROM,  UASM,  USNM. 

TYPE-LOCALITY.— Hoback  River  (8  miles  [ca.  13  km]  NW 
of  Bondurant),  Sublette  County,  Wyoming. 


DIAGNOSTIC  COMBINATION. — Figure  1;  size 
very  large,  standardized  body  length  male  great- 
er than  12.0  mm,  female  greater  than  12.5  mm; 
head  moderate  in  size  and  width,  dark,  with  a 
pair  of  pale  paramedial  spots  on  vertex;  prono- 
tum (Fig.  27)  slightly  flattened,  only  slightly  cor- 
date, broad  basally,  midlateral  seta  present,  ba- 
solateral seta  present;  elytra  with  marked  violet, 
blue,  or  green  metallic  reflection,  elytral  silhou- 
ette subrectangular,  broad  basally;  hind  coxa 
(Fig.  36)  with  four  to  six  basal  and  one  (in  very 
few  individuals  two  or  three)  apical  seta(-ae); 
specimen  from  locality  in  western  Wyoming 
(Fig.  69). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  subspecies  in  honor  of 
my  wife,  Beverly  Ann  Kavanaugh,  in  grateful 
acknowledgment  of  her  constant  support  through 
the  course  of  this  study. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  69; 
at  present  known  only  from  Hoback  River  at  the 
south  slope  of  the  Gros  Ventre  Mountains.  I 
have  seen  specimens  from  the  following  locality. 

United  States  of  America 

WYOMING:  Sublette  County,  Bondurant  (8  miles  [ca.  13  km] 
NW  at  Hoback  River  [2,100  m])  [July-Aug.]  (255;  CAS, 
DHKa,  FMNH,  MCZ,  UASM,  USNM). 

Specimens  Without  Locality  Data:  (1:  ROM). 

Nebria  trifaria  utahensis,  new  subspecies 

(Figures  8,  28,  45,  50,  70) 

HOLOTYPE,  a  male,  in  CNC,  labelled:  "Lonesome  Beaver, 
7500',  Henry  Mts.,  Utah  VII  20-22,  68  H.  F.  Howden"/ 
"Holotype  Nebria  trifaria  utahensis  Kavanaugh  det.  D.  H. 
Kavanaugh  1976"  [red  label].  ALLOTYPE  (same  data  as  holo- 
type) also  in  CNC.  In  total,  13  PARATYPES  (9  males  and  4 
females)  are  deposited  in  the  following  collections:  CAS, 
CNC,  UASM. 

TYPE-LOCALITY. — Lonesome  Beaver,  Henry  Mountains, 
Garfield  County,  Utah. 

DIAGNOSTIC  COMBINATION. — Head  moderate 
in  size  and  width,  dark,  with  a  pair  of  pale  par- 
amedial spots  on  vertex;  antennal  scape  (Fig.  8) 
slightly  arcuate  or  straight,  slightly  to  markedly 
narrowed  basally;  pronotum  (Fig.  28)  with  basal 
sinuation  of  lateral  margin  moderately  deep, 
midlateral  seta  present,  basolateral  seta  present; 
elytra  without  metallic  reflection,  microsculp- 
ture  deeply  impressed,  meshes  isodiametric, 
elytral  silhouette  subovoid;  hind  wing  reduced  in 
width  and  length,  truncate  distal  to  stigma;  male 
with  apex  of  median  lobe  as  in  Figure  45;  female 
with  posterodorsal  sclerite  in  bursa  copulatrix 


KAVANAUGH:  NEW  NEARCTIC  NEBR1A 


111 


as  in  Figure  50;  specimen  from  Henry  Moun- 
tains (Utah)  (Fig.  70). 

DERIVATION  OF  TAXON  NAME. — This  subspe- 
cies is  named  for  the  State  of  Utah,  in  which  its 
geographical  range  is  restricted. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  70; 
restricted  to  the  Henry  Mountains  of  southcen- 
tral  Utah.  I  have  studied  specimens  from  the 
following  localities. 

United  States  of  America 

UTAH:  Garfield  County,  Hanksville  (24  miles  [ca.  39  km|  s 
[2,290  m])  [July]  (3;  CNC),  Lonesome  Beaver  ([2,290  m]) 
[July]  (12:  CAS,  CNC,  UASM). 

NOMENCLATURAL  NOTES  AND  LECTOTYPE 
DESIGNATIONS 

Carabus  gyllenhali  SCHONHERR,  1806:196. 
[=Nebria  gyllenhali  (Schonherr).]  LECTO- 
TYPE  (here  designated),  a  male,  in  NRSS,  la- 
belled: "cTY  [triangular  blue  label]/  "Lecto- 
type  Carabus  gyllenhali  Schonherr  designated 
by  D.  H.  Kavanaugh  1976"  [red  label].  Five 
paralectotypes  (same  data  as  lectotype)  also 
in  NRSS. 

The  name  Carabus  rufescens  Str0m  (1768:331) 
did  not  appear  in  the  literature  between  the  date 
of  its  original  description  and  1880,  when  Sch0y- 
en  (1880:183)  revived  its  use  (Carl  Lindroth,  per- 
sonal communication).  Jeannel  (1937:4),  how- 
ever, accepted  N.  rufescens  as  the  senior 
synonym  of  N.  gyllenhali;  and,  following  him, 
a  number  of  authors  have  subsequently  applied 
the  name  "rufescens"  to  this  species  (Blair 
1950:220;  Friden  1971:30;  Greenslade  1968:41; 
Horvatovitch  1973:252;  Luff  1972:174;  Luff  and 
Davies  1972:47;  Nakane  1963:18,  1971:180;  and 
Ueno  1953:59,  1955:47).  The  combination  A/. 
rufescens  gyllenhali  has  sporadically  appeared 
(Jeanne  1966:12;  and  Jeannel  1937:4,  1941:19)  as 
has  the  unfortunate  combination  N.  gyllenhali 
rufescens,  where  "rufescens"  referred  various- 
ly to  a  subspecies,  race,  variety,  or  aberration 
of  N.  gyllenhali  (Andrewes  1939:161;  Banninger 
1925:345,  1960:261;  Csiki  1927:359;  Munster 
1933:267;  and  Reitter  1908:93). 

The  name  given  to  this  particular  species  is  of 
considerable  importance.  The  accumulated  lit- 
erature on  this  species  is  very  extensive,  and  its 
members  are  among  the  most  broadly  distrib- 
uted and  commonly  collected  of  all  carabid  bee- 
tles. Both  names  have  been  so  often  used  in  the 
literature  that  a  decision  for  or  against  either  will 


cause  some  confusion,  but  it  is  still  more  im- 
portant to  reach  some  decision  which  will  pro- 
vide for  consistency  in  all  future  usage.  Lindroth 
(1939:59,  1954:121,  1961:78)  and  Banninger 
(1949:144)  have  refused  to  recognize  N.  rufes- 
cens as  the  valid  name  for  this  species.  For  the 
reasons  they  have  cited,  because  no  type-spec- 
imen has  been  found  or  perhaps  ever  existed, 
and  because  Str0m's  original  description  "can- 
not be  interpreted  (not  even  as  a  Nebria)"  (Carl 
Lindroth,  personal  communication),  I  concur 
that  the  name  N.  gyllenhali  (Schonherr)  should 
be  conserved  as  the  valid  name  for  the  species. 
Any  future  use  of  the  epithet  rufescens  should 
be  discouraged;  and  the  name  Carabus  rufes- 
cens Str0m  should  be  listed  as  a  nomen  dubium. 

Carabus  nivalis  PAYKULL,  1790:52.  [=Nebria 
nivalis  (Paykull).]  LECTOTYPE  (here  designat- 
ed), a  male,  in  NRSS,  labelled:  "Mus. 
Payk.'V  "Riksmuseum  Stockholm"  [green  la- 
bel]/ "Lectotype  Carabus  nivalis  Paykull  des- 
ignated by  D.  H.  Kavanaugh  1976"  [red  la- 
bel]. 

The  specimen  chosen  as  lectotype  of  Carabus 
nivalis  is  one  of  three  specimens  so  identified  in 
NRSS  from  the  Paykull  Collection.  The  other 
two  specimens  are  actually  Nebria  gyllenhali 
gyllenhali  (Schonherr)  adults.  Because  all  three 
specimens  bear  identical  labels  and  have  tradi- 
tionally been  considered  syntypes,  the  two  N. 
gyllenhali  specimens  could  perhaps  be  consid- 
ered paralectotypes.  Paykull  (1798:119)  subse- 
quently described  a  "variety  B"  of  C.  nivalis, 
which  is  clearly  synonymous  with  N.  gyllenhali 
gyllenhali  and  was  based  on  specimens  from  a 
different  locality  and  collector.  The  original  de- 
scription of  C.  nivalis  does  not  fit  the  two  het- 
ero-specific  specimens,  but  the  description  of 
"variety  B"  does  so.  I  conclude  that  these  spec- 
imens are  probably  not  part  of  Paykull's  original 
type-series  of  C.  nivalis;  but  they  may  be  the 
specimens  he  described  later  as  "variety  B." 

Helobia  castanipes  KIRBY,  1837:20.  [=Nebria 
gyllenhali  castanipes  (Kirby).]  LECTOTYPE 
(here  designated),  a  female,  in  BMNH,  la- 
belled: "Type"  [red-trimmed  disk]/  "70' V 
"N.  Amer."  [pale  blue  disk  with  "5699"  on 
underside]/  "apparently  Type  of  castanipes 
Kby.  Lindroth  1952"/  "Lectotype  Helobia 
castanipes  Kirby  designated  by  D.  H.  Kavan- 
augh 1976"  [red  label]. 


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The  lectotype  bears  no  label  in  Kirby's  hand. 
However,  according  to  P.  M.  Hammond  (per- 
sonal communication),  the  number  "5699"  on 
the  underside  of  one  label  refers  to  an  entry  in 
manuscript  catalogs  of  the  Coleoptera  in 
BMNH.  An  original  label  in  Kirby's  hand  is 
pasted  in  the  catalog  at  this  entry  and  reads  "73. 
1.  9?/  castanipes."  The  fate  of  the  second  of 
two  specimens  mentioned  by  Kirby  in  his  orig- 
inal description  is  unknown. 

Nebria  arkansana  CASEY,  1913:52. 

Lindroth  (1961:70)  was  certainly  correct  in 
denying  the  validity  of  Casey's  records  for  Ar- 
kansas and  Indiana  for  this  species;  and  his 
choice  for  the  new  type-locality  ("Valley  of  the 
Upper  San  Juan  River/'  Archuleta  County, 
"Colorado")  is  quite  reasonable.  The  specimens 
in  Casey's  series  are  labelled  "Ind"  and  "S. 
Ark";  and  it  is  at  least  possible  that  the  latter 
refers  to  the  South  Arkansas  River  (a  quite  suit- 
able locality  for  the  species,  in  Chaffee  County, 
Colorado)  rather  than  to  the  State  of  Arkansas 
as  supposed  by  Casey.  It  is  also  possible  that 
"Ind"  refers  to  the  "Indian  Territory,"  al- 
though general  use  of  that  name  for  the  region 
was  abandoned  long  before  1913. 

Nebria  bifaria  MANNERHEIM,  1853:120.  [=Ne- 
bria  nivalis  nivalis  (Paykull).]  LECTOTYPE 
(here  designated),  a  male,  in  ZILR,  labelled: 
"38"/  "Nebria  carbonaria  Esch.  Insel.  Paul"/ 
"Lectotype  ?"  [red  label]/  "Lectotype  Nebria 
bifaria  Mannerheim  designated  by  D.  H.  Ka- 
vanaugh  1976"  [red  label]. 

Nebria  catenata  CASEY,  1913:49.  [=Nebria  tri- 
faria  catenata  Casey.]  TYPE-LOCALITY. — 
Colorado;  restricted  by  Erwin  and  Ball 
(1972:97)  to  San  Juan  Mountains,  and  here 
further  restricted  to  Wolf  Creek  Pass,  Mineral 
County,  Colorado. 

Nebria  diversa  LECONTE,  1863a:2  [as  a  replace- 
ment name  for  N.  livida  LeConte].  LECTO- 
TYPE (same  as  for  N.  livida  LeConte;  see  be- 
low for  data). 

Nebria  elias  MOTSCHULSKY,  1865:274.  The  type- 
specimen  of  N.  elias  has  been  completely  de- 
stroyed and  only  its  pin  and  labels  remain  (in 
ZMUM).  Motschulsky's  original  description 
is  too  vague  to  permit  assurance  that  N.  elias 
and  N.  gyllenhali  castanipes  are  synony- 


mous. The  occurrence  of  the  latter  in  Alaska 
(the  type  area  of  N.  elias)  is  restricted  and 
local,  which  casts  further  uncertainty  as  to 
appropriate  application  of  the  name.  I  there- 
fore refrain  from  designation  of  a  neotype  and 
list  N.  elias  as  a  nomen  dubium,  but,  for  con- 
venience, place  it  as  a  synonym  of  N.  gyllen- 
hali castanipes,  following  Lindroth  (1961:78). 

Nebria  eschscholtzii  MENETRIES,  1844:55.  LEC- 
TOTYPE (here  designated),  a  female,  in  ZILR, 
labelled:  [gold-coated  square]/  "California" 
[pink  label]/  "Eschscholtzii  Menet.  Californ."/ 
"Syntypus"  [red  label]/  "Lectotype  Nebria 
eschscholtzii  Menetries  designated  by  D.  H. 
Kavanaugh  1976"  [red  label].  One  paralecto- 
type  also  in  ZILR.  TYPE-LOCALITY. — Califor- 
nia; here  restricted  to  South  Fork  of  American 
River,  3  miles  [ca.  4.8  km]  w  of  Riverton,  El 
Dorado  County,  California. 

Nebria  expansa  CASEY,  1913:56.  [=Nebria  la- 
custris  Casey,  1913:56.]  TYPE-LOCALITY. — 
Indiana;  here  restricted  to  Turkey  Run  State 
Park,  Parke  County,  Indiana. 

The  name  N.  lacustris  Casey  has  priority  over 
N.  expansa  by  precedence  of  position  as  well 
as  by  action  of  first  reviser  (Bell  1955:265;  see 
also  Lindroth  1961:77).  In  his  original  descrip- 
tion, Casey  recorded  N.  expansa  from  "Texas 
and  Indiana,"  based  on  his  study  of  two  fe- 
males. Bell  (1955:267)  and  Lindroth  (1961:77, 
1975:112)  accepted  the  record  from  Texas.  In 
my  judgment,  this  record  must  be  erroneous. 
Acting  on  my  suggestion,  Lindroth  (1975:147) 
amended  his  lectotype  designation  (1975:112) 
and  selected  the  Casey  specimen  labelled  "L.," 
interpreted  by  Casey  as  from  Indiana,  instead  of 
the  "Texas"  specimen.  Individuals  recognizable 
as  the  "expansa"  form  of  N.  lacustris  have 
often  been  collected  at  the  restricted  type-local- 
ity chosen. 

Nebria  fusiformis  VAN  DYKE,  1926: 11.  [=Nebria 
spatulata  spatulata  Van  Dyke.] 

Because  this  name  appeared  unaccompanied 
by  any  description  or  indication,  it  should  be 
considered  a  nomen  nudum.  It  is  clear  to  me, 
however,  from  the  text  in  which  the  name  ap- 
peared, that  Van  Dyke  was  referring  to  his  N. 
spatulata,  the  original  description  of  which  ap- 
peared the  previous  year. 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


113 


Nebria  gebleri  DEJEAN,  1831:573.  HOLOTYPE,  a 
female,  in  MHNP,  labelled:  "Sitka"/  "Ge- 
bleri Eschs"/  "Ex  Musaeo  Mniszech'V  "Ho- 
lotype  Nebria  gebleri  Dejean  del.  D.  H.  Ka- 
vanaugh  1976"  [red  label]. 

It  is  clear  from  his  original  description  that  the 
specimen  of  N.  gebleri  seen  by  Dejean  was  at 
that  time  complete.  In  his  review  of  the  Dejean 
types,  Lindroth  (1955b:12)  noted  that  the  head 
and  prothorax  of  the  type-specimen  had  been 
lost.  In  my  study  of  the  type  (in  1976),  I  found 
it  to  be  complete  again!  Unfortunately,  the  parts 
replaced  are  not  only  from  a  different  specimen 
but  also  from  one  representing  a  different 
species  [namely,  Nebria  pic icornis  (Fabricius)]. 
There  is  no  doubt,  however,  that  the  pterothor- 
ax  and  abdomen  (through  which  the  pin  passes) 
are  the  recognizable  remains  of  the  valid  type- 
specimen. 

Nebria  gregaria  FISCHER  VON  WALDHEIM, 
1821:72.  LECTOTYPE  (here  designated),  a 
male,  in  ZMUM,  labelled:  "47.  gregaria 
Fisch.'V  "Lectotype  Nebria  gregaria  Fischer 
v.  Wald.  designated  by  D.  H.  Kavanaugh 
1976"  [red  label]. 

Fischer  (1821:73)  indicated  that  his  descrip- 
tion of  N.  gregaria  was  based  on  material  in  his 
own  and  Eschscholtz's  collections.  I  have  been 
unable  to  locate  any  Eschscholtz  specimens  of 
N.  gregaria  from  among  those  in  UMHF.  Ap- 
parently, the  single  Fischer  specimen  (ZMUM) 
chosen  as  lectotype  is  the  only  survivor  from 
the  type-series. 

Nebria  hudsonica  LECONTE,  1863b:3.  LECTO- 
TYPE (here  designated),  a  male,  in  MCZ,  la- 
belled: "Saskatchewan"/  "Type  643"  [red  la- 
bel]/ "N.  hudsonica  LeC. "I  "Lectotype 
Nebria  hudsonica  LeConte  designated  by  D. 
H.  Kavanaugh  1976"  [red  label].  TYPE-LO- 
CALITY.— Saskatchewan,  Hudson's  Bay  Ter- 
ritory; here  restricted  to  North  Saskatchewan 
River  at  Rocky  Mountain  House,  Alberta. 

In  1863,  the  area  now  occupied  by  the  Prov- 
ince of  Saskatchewan  was  part  of  Hudson's  Bay 
Territory  (or  the  Northwest  Territories),  and  at 
least  up  to  that  date,  the  name  "Saskatchewan" 
was  apparently  applied  only  to  the  Saskatche- 
wan River  system.  Therefore,  LeConte's  use  of 
the  name  as  a  type-locality  probably  refers  to 
the  river  system  rather  than  the  province.  Only 
those  parts  of  the  system  in  Alberta  are  within 


the  continuous  range  of  N ' .  hudsonica,  although 
I  have  seen  two  specimens  from  localities  in  the 
Province  of  Saskatchewan.  My  selection  of  the 
restricted  type-locality  reflects  these  facts  and 
considerations. 

Nebria  incerta  CASEY,  1913:53.  [= Nebria  obli- 
qua  LeConte.]  TYPE-LOCALITY. — Colorado: 
here  restricted  to  North  Fork  of  South  Platte 
Canyon  at  Santa  Maria,  Park  County,  Colo- 
rado. This  restricted  type-locality  is  the  same 
as  for  N.  obliqua. 

Nebria  ingens  HORN,  1870:98.  LECTOTYPE  (here 
designated),  a  female,  in  MCZ,  labelled: 
"Gala."/  "Type  No.  1026"  [red  label]/  "Ne- 
bria ingens  Horn"/  "Lectotype  Nebria  ingens 
Horn  designated  by  D.  H.  Kavanaugh  1976" 
[red  label].  One  female  paralectotype,  also  in 
MCZ,  labelled:  "Gala."/  "Type  8127  [red  la- 
bel]/ "N.  ingens  Horn."  TYPE-LOCALITY. — 
Sierra  Nevada  Mountains  east  of  Visalia,  Cal- 
ifornia; here  restricted  to  Franklin  Lakes,  Tu- 
lare  County,  California. 

In  his  original  description,  Horn  mentioned 
two  specimens,  a  male  and  a  female.  The  MCZ 
specimen  seen  by  Lindroth  (1961:87),  his  "al- 
lotype,"  is  a  female.  The  other  specimen,  from 
the  Horn  Collection  (formerly  at  ANSP,  now  at 
MCZ),  is  also  a  female;  so  Horn  did  not  actually 
see  a  male  of  this  species.  The  lectotype  chosen 
is  the  specimen  from  the  Horn  Collection. 

Nebria  livida  LECONTE,  1859:84  [preoccupied 
by  Carabus  lividus  Linnaeus,  1758:414;  =Ne- 
bria  diversa  LeConte].  LECTOTYPE  (here  des- 
ignated), a  male,  in  MCZ,  labelled:  [blue  disk]/ 
"Type  642"  [red  label]/  "N.  diversa  LeC.  \\ 
livida  LeC."/  "Lectotype  Nebria  livida 
LeConte  designated  by  D.  H.  Kavanaugh 
1976"  [red  label].  One  male  paralectotype 
(same  data  as  lectotype)  also  in  MCZ. 

Nebria  longula  LECONTE,  1878:478.  [=Nebria 
suturalis  LeConte — NEW  SYNONYMY.]  TYPE- 
LOCALITY. — Colorado;  here  restricted  to 
Longs  Peak,  Rocky  Mountain  National  Park, 
Colorado. 

To  date,  all  Colorado  records  for  N.  suturalis 
have  been  from  the  Front  Range  of  the  Rocky 
Mountains.  The  restricted  type-locality  is  a  pro- 
tected wilderness  area  in  that  range. 


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Nebria  mannerheimii  FISCHER  VON  WALDHEIM, 
1828:253.  LECTOTYPE  (here  designated),  a 
male,  in  ZMUM,  labelled:  "Sitcha"/  "48. 
mannerheimii  Fisch."/  "Lectotype  Nebria 
mannerheimii  Fischer  v.  Wald.  designated  by 
D.  H.  Kavanaugh  1976"  [red  label].  One  male 
paralectotype  in  UMHF. 

Both  specimens  that  I  recognize  as  comprising 
the  type-series  of  N.  mannerheimii  were  prob- 
ably those  collected  by  Eschscholtz  (Fischer 
von  Waldheim  1828:253).  The  specimen  chosen 
as  lectotype  is  from  the  Fischer  Collection 
(ZMUM).  The  paralectotype  is  from  the  Man- 
nerheim  Collection  (UMHF)  and  bears  an 
"Eschsch."  label. 

Nebria  metallica  FISCHER  VON  WALDHEIM, 
1821:71.  LECTOTYPE  (here  designated),  a 
male,  in  ZMUM,  labelled:  "Unalaschka.  F."/ 
"73  metallica  Fisch."/  "Lectotype  Nebria 
metallica  Fischer  v.  Wald.  designated  by  D. 
H.  Kavanaugh  1976"  [red  label].  One  male 
and  one  female  paralectotype  are  in  UMHF. 

Fischer's  description  of  N.  metallica  was 
based  on  material  in  his  own  and  Eschscholtz's 
collections  (Fischer  von  Waldheim  1821:72). 
Specimens  from  the  latter,  identified  as  such, 
are  now  in  UMHF  as  part  of  the  Mannerheim 
Collection.  I  have  chosen  the  Fischer  Collection 
specimen  as  lectotype,  the  two  Eschscholtz 
specimens  as  paralectotypes.  A  fourth  speci- 
men, in  the  Fischer  Collection  (ZMUM),  bears 
a  "Kadjak"  [=Kodiak]  label  and  is  thus  exclud- 
ed from  the  type-series. 

Nebria  moesta  LECONTE,  1850:209.  [=Nebria 
gyllenhali  castanipes  (Kirby).]  LECTOTYPE 
(here  designated),  a  male,  in  MCZ,  labelled: 
[light  green  disk]/  "Type  645"  [red  label]/  "N. 
moesta  LeC."/  "N.  castanipes  Kby."/  "Lec- 
totype Nebria  moesta  LeConte  designated  by 
D.  H.  Kavanaugh  1976"  [red  label].  One  male 
and  two  female  paralectotypes  (same  data  as 
lectotype)  also  in  MCZ. 

Nebria  mollis  MOTSCHULSKY,  1865:274.  [=Ne- 
bria  nivalis  nivalis  (Paykull).]  LECTOTYPE 
(here  designated),  sex  undetermined  (speci- 
men incomplete,  badly  damaged),  in  ZMUM, 
labelled:  "A.  borealis"  [green  label]/  "Nebria 
mollis  Motsch.  Am.  arc.  cont."  [green  label]/ 
[red  rectangle]/  "Lectotype  Nebria  mollis 


Motschulsky  designated  by  D.  H.  Kavanaugh 
1976"  [red  label]. 

In  his  original  description,  Motschulsky  pre- 
sented the  species  epithet  as  "molbis."  I  have 
found  no  published  statement  by  Motschulsky 
that  "molbis"  was  a  misspelling.  However,  the 
type-specimen  is  labelled  "mollis";  from  as  ear- 
ly as  1868,  every  subsequent  citation  of  the 
name  has  used  the  latter  spelling.  This  suggests 
early  recognition,  possibly  initiated  (privately) 
by  Motschulsky  himself,  of  an  original  error. 
The  first  clear  indication  of  an  intended  emen- 
dation is  that  of  Csiki  (1927:389). 

Nebria  miihlenbergii  STURM,  1826:173  [here 
emended  to  Nebria  muehlenbergii  Sturm]. 

Because  Sturm  never  selected  type-speci- 
mens, none  exists  for  this  name.  Furthermore, 
no  description  ever  accompanied  the  use  of  this 
name.  However,  Sturm  (1843:16)  subsequently 
cited  N.  muehlenbergii  as  a  synonym  of  N.  pal- 
lipes,  thereby  providing  the  only  clue  to  his  ap- 
plication of  the  name  to  a  taxon.  The  name 
should  be  treated  as  a  nomen  nudum. 

Nebria  obliqua  LECONTE,  1866c:363.  LECTO- 
TYPE (here  designated),  a  male,  in  MCZ,  la- 
belled: "Col."/  "Type  646"  [red  label]/  "Ne- 
bria obliqua  Lee.  Lewis  Col."/  "obliqua  2"/ 
"Lectotype  Nebria  obliqua  LeConte  desig- 
nated by  D.  H.  Kavanaugh  1976"  [red  label]. 
One  female  paralectotype  (same  data  as  lec- 
totype) also  in  MCZ.  TYPE-LOCALITY. — Col- 
orado; here  restricted  to  North  Fork  of  South 
Platte  Canyon  at  Santa  Maria,  Park  County, 
Colorado. 

The  type-locality  of  N.  obliqua,  as  originally 
stated,  was  Colorado.  LeConte  (1878:478)  sub- 
sequently listed  "North  Fork  of  South  Platte 
Canon  (7,000  to  8,000  ft.)"  as  a  locality  for  the 
species,  and  Lindroth  (1961:73)  accepted  this  as 
the  type-locality.  I  here  formally  restrict  the 
type-locality  to  that  area. 

Nebria  obtusa  LECONTE,  1878:478.  [=Nebria 
obliqua  LeConte — NEW  SYNONYMY.] 

My  interpretation  of  LeConte' s  original  de- 
scription of  N.  obtusa  is  that  the  name  is  based 
on  a  single  specimen  (see  LeConte  1878:479, 
lines  7-8).  I  therefore  consider  the  type-speci- 
men (in  MCZ)  to  be  a  holotype.  The  synonymy 
noted  here  was  actually  suggested  by  LeConte 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


115 


(1878:479)  and  is  supported  by  abundant  data 
(Kavanaugh,  manuscript  in  preparation). 

Nebria  oregona  CASEY,  1913:52.  [=Nebria  man- 
nerheimii  Fischer  von  Waldheim.]  TYPE-LO- 
CALITY.— Clackamas  County,  Oregon;  here 
restricted  to  Zigzag  River  at  Rhododendron, 
Clackamas  County,  Oregon. 

Nebria  ovipennis  LECONTE,  1878:477.  TYPE-LO- 
CALITY.— Sierra  Nevada,  California;  here  re- 
stricted to  Mount  Conness  (east  slope,  above 
Greenstone  Lake),  Mono  County,  California. 

The  holotype  of  N.  ovipennis  (in  MCZ)  is  rep- 
resentative of  populations  north  of  Mount  Lyell, 
Yosemite  National  Park,  but  not  south  of  that 
point  in  the  Sierra  Nevada.  The  restricted  type- 
locality  chosen  is  therefore  both  more  precise 
and  appropriate  for  the  form  represented  by  the 
type-specimen. 

Nebria  pallipes  SAY,  1823:78. 

Lindroth  (1969a:1149,  and  in  Lindroth  and 
Freitag  1969:326)  discussed  the  problem  of  es- 
tablishing the  actual  publication  date  of  Say's 
paper  in  which  the  original  description  of  N. 
pallipes  appeared.  His  comments,  however, 
support  the  date  cited  above,  rather  than  1825, 
the  year  Lindroth  (1961:76)  cited. 

Nebria  rathvoni  LECONTE,  1853:400.  [=Nebria 
gebleri  rathvoni  LeConte — NEW  STATUS.] 
HOLOTYPE,  a  male,  in  MCZ,  labelled:  "6456 
ft.  Lake  Tahoe,  Cal.  May  24,  1879'V  [light 
green  square]/  "79."  [red-tipped  label]/  "N. 
rathvoni  LeC.'V  "Type  7403"  [red  label]/ 
"Holotype  Nebria  rathvoni  LeConte  det.  D. 
H.  Kavanaugh  1976"  [red  label].  TYPE-LO- 
CALITY.— Sacramento,  California;  here 
emended  to  Truckee  River  at  Truckee,  Ne- 
vada County,  California. 

There  is  no  reason  to  doubt  that  the  specimen 
recognized  here  as  the  holotype  of  N.  rathvoni 
is  actually  the  specimen  seen  by  LeConte,  al- 
though locality  and  date  data  on  one  label  indi- 
cate otherwise.  It  is  the  only  specimen  of  this 
taxon  in  the  LeConte  Collection  (MCZ).  Fur- 
thermore, the  troublesome  specimen  label  itself 
appears  to  be  relatively  new,  and  I  suspect  that 
it  has  been  more  recently  added  to  the  original 
type-specimen  and  thereby  represents  a  misla- 
belling. 


LeConte's  type-locality,  Sacramento,  is  well 
outside  the  habitat  range  of  the  taxon;  so  the 
type  was  surely  not  collected  there  (although 
Sacramento  was  probably  the  closest  important 
town  to  the  true  collection  site.  Truckee,  the 
emended  type-locality,  is  about  at  the  midpoint 
of  both  the  habitat  and  geographical  ranges  of 
the  taxon. 

Nebria  sahlbergii  FISCHER  VON  WALDHEIM, 
1828:254.  LECTOTYPE  (here  designated),  a 
male,  in  UMHF,  labelled:  "d"/  "Eschsch."/ 
"Sitka'V  "Lectotype  Nebria  sahlbergii  Fi- 
scher v.  Wald.  designated  by  D.  H.  Kavan- 
augh 1976"  [red  label]/  "Lectotype  Nebria 
violacea  Motsch.  designated  by  D.  H.  Kavan- 
augh 1976"  [red  label]. 

Fischer's  description  of  N.  sahlbergii  was 
based  on  a  study  of  Eschscholtz  material  (Fi- 
scher von  Waldheim  1828:254).  I  have  located 
only  two  specimens  probably  representing  the 
original  type-series — one  in  the  Fischer  Collec- 
tion (ZMUM),  the  other  in  the  Mannerheim  Col- 
lection (UMHF).  I  have  chosen  the  UMHF 
specimen  as  lectotype  because,  first,  the  original 
description  better  fits  this  specimen,  and,  sec- 
ond, the  Fischer  (ZMUM)  specimen  is  actually 
a  representative  of  N.  gyllenhali  castanipes 
rather  than  N.  sahlbergii  as  the  latter  name  was 
applied  by  Lindroth  (1961:68).  The  Fischer 
specimen  must  be  considered  a  paralectotype  of 
N.  sahlbergii  in  spite  of  its  identity.  In  fact,  the 
strange  misuse  of  the  name  "sahlbergii"  by  nu- 
merous North  American  authors  in  referring  to 
N.  gyllenhali  castanipes  (Kirby)  (e.g.,  Casey 
1913:51;  Hatch  1939:120;  Horn  1870:102;  Le- 
Conte 1853:400,  1878:479)  can  only  now  be  ex- 
plained— that  is,  if  these  authors  assumed  that 
the  Fischer  specimen  was  the  type  of  N.  sahl- 
bergii. I  choose  not  to  follow  their  interpretation 
in  order  to  preserve  the  name  Helobia  castan- 
ipes Kirby  (see  above),  which  would  become  an 
objective  junior  synonym  of  N .  sahlbergii  if  the 
Fischer  specimen  were  designated  as  lectotype. 

Nebria  suturalis  LECONTE,  1850:209.  LECTO- 
TYPE (here  designated),  a  male,  in  MCZ,  la- 
belled: [light  green  disk]/  "Type  650"  [red  la- 
bel]/ "suturalis  3"/  "Lectotype  Nebria 
suturalis  LeConte  designated  by  D.  H.  Ka- 
vanaugh 1976"  [red  label].  Two  female  para- 
lectotypes  (same  data  as  lectotype)  also  in 
MCZ. 


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Nebria  tenuipes  CASEY,  1913:51.  [=Nebria  esch- 
scholtzii  Menetries.]  TYPE-LOCALITY. — Ala- 
meda  County,  California;  here  restricted  to 
Niles  Canyon,  Alameda  County,  California. 

Nebria  texana  CASEY,  1913:54.  [=Nebria  obli- 
qua  LeConte.]  TYPE-LOCALITY. — "Texas." 

The  holotype  of  N.  texana  (in  USNM)  bears 
the  label  "Tex,"  interpreted  by  Casey  as  refer- 
ring to  the  State  of  Texas  (a  highly  improbable 
record).  It  is  more  probable  that  the  specimen 
was  collected  in  Colorado,  perhaps  at  Texas 
Creek,  a  small  tributary  of  the  Arkansas  River, 
Fremont  County,  Colorado.  However,  I  prefer 
not  to  amend  or  restrict  the  type-locality  at  pres- 
ent, pending  further  field  search  in  the  moun- 
tainous regions  of  western  Texas. 

Nebria  trifaria  LECONTE,  1878:478.  LECTOTYPE 
(here  designated),  a  female,  in  MCZ,  labelled: 
"9500  ft.  Amer.  Fork  Canon  Utah,  Aug.  2-3, 
1877'V  "Type  651"  [red  label]/  "N.  trifaria 
LeC."l  "Lectotype  Nebria  trifaria  LeConte 
designated  by  D.  H.  Kavanaugh  1976"  [red 
label]. 

Nebria  vandykei  BANNINGER,  1928:5.  LECTO- 
TYPE (here  designated),  a  male,  in  ETHZ,  la- 
belled: "Paradise  Val.  Mt.  Rainier  Wash.  VII- 
1 8-1 920' V  "Col.  by  E.  C.  Van  Dyke"/  "1. 
Fuhl.gld.  as.  1+2  B."/  "FG.  Basig.  bil. 
OP!"/  "Nebria  trifaria  LeC."/  "Type  Nebria 
Van  Dykei  10.1927"  [red-trimmed  label]/ 
"Lectotype  Nebria  vandykei  Banninger  des- 
ignated by  D.  H.  Kavanaugh  1976"  [red  la- 
bel]. Two  paralectotypes  also  in  ETHZ. 

In  his  original  description,  Banninger  men- 
tioned four  specimens  in  the  type-series,  but 
only  three  of  these  are  accounted  for  at  present 
(W.  Sauter,  personal  communication). 

Nebria  violacea  MOTSCHULSKY,  1850:73.  [=Ne- 
bria  sahlbergii  sahlbergii  Fischer  von  Wal- 
dheim.]  LECTOTYPE  (here  designated),  same 
specimen  as  lectotype  of  Nebria  sahlbergii 
Fischer  von  Waldheim  (see  above). 

Mannerheim  apparently  obtained  and  exam- 
ined the  Eschscholtz  specimen  of  N.  sahlbergii 
now  in  UMHF,  studied  the  specimen  identified 
as  N.  sahlbergii  in  Fischer's  Collection  (ZMUM), 
and  then  noted  (Mannerheim  1843:189)  that  two 
"varieties"  existed.  He  called  the  form  repre- 
sented by  the  Eschscholtz  specimen  in  his  pos- 


session "var.  B."  Motschulsky  (1850:73),  refer- 
ring to  Mannerheim's  brief  description  of  variety 
"B,"  named  this  form  N.  violacea.  Conse- 
quently, the  specimen  now  in  UMHF  appears 
to  be  the  specimen  upon  which  both  N.  violacea 
and  N.  sahlbergii  are  based.  I  have  therefore 
designated  and  so  labelled  the  specimen  as  lec- 
totype for  both  names. 

Nebria  viridis  HORN,  1870:101.  [=Nebriafrigida 
R.  F.  Sahlberg.]  LECTOTYPE  (here  designat- 
ed), a  male,  in  MCZ,  labelled:  "R.  [super- 
script "M"]  A."/  "Type  No.  1027-"  [red  la- 
bel]/ "N.  viridis  Horn"/  "Lectotype  Nebria 
viridis  Horn  designated  by  D.  H.  Kavanaugh 
1976"  [red  label].  One  male  paralectotype 
(same  data  as  lectotype)  also  in  MCZ. 

In  Horn's  original  description,  only  two  spec- 
imens are  mentioned.  There  are  now  three  spec- 
imens in  MCZ  (one  from  the  Horn  Collection, 
two  from  the  LeConte  Collection).  I  have  des- 
ignated the  Horn  specimen  as  lectotype  and  the 
first  LeConte  specimen  as  the  paralectotype. 
The  second  LeConte  specimen  (labelled  "viridis 
2")  has  no  type  status.  All  three  specimens  bear 
identical  first  labels;  but  LeConte  may  have  re- 
ceived his  second  specimen  directly  from  Ulke 
(see  Horn  1870:14),  his  first  specimen  from 
Horn. 

ACKNOWLEDGMENTS 

I  gratefully  acknowledge  the  assistance  of 
those  curators  and  other  individuals  listed  above 
(under  Materials  and  Methods)  who  loaned  ma- 
terial in  their  care  to  me  for  study.  In  addition, 
the  following  individuals  made  type-material 
available  to  me  on  loan  or  provided  critical  in- 
formation on  type-specimens:  T.  L.  Erwin 
(USNM),  P.  M.  Hammond  (BMNH),  H.  Hippa 
(UMTF),  S.  Keleinikova  (ZMUM),  O.  L.  Kry- 
zhanovskij  and  V.  G.  Schilenkov  (ZILR),  J.  F. 
Lawrence  and  J.  Scott  (MCZ),  H.  B.  Leech 
(CAS),  O.  Martin  (ZMKD),  T.  Nyholm  (NRSS), 
H.  Perrin  (MHNP),  S.  Rohwer  (UWBM),  W. 
Sauter  (ETHZ),  G.  Scherer  (ZSBS),  and  H. 
Silfverberg  (UMHF).  T.  L.  Erwin  (USNM),  M. 
H.  Hatch  (UWBM),  J.  F.  Lawrence  (MCZ),  H. 
B.  Leech  (CAS),  P.  Oman  (OSUO),  L.  L.  Pe- 
chuman  (CUIC),  A.  Provonsha  (PURC),  and  A. 
Smetana  (CNC)  were  helpful  hosts  during  my 
visits  to  study  material  in  their  respective  insti- 
tutions. I  thank  also  the  many  colleagues  and 


KAVANAUGH:  NEW  NEARCTIC  NEBRIA 


117 


friends  who  assisted  me  in  the  field  by  providing 
companionship,  physical  labor,  and  other  re- 
sources in  support  of  the  project,  and  my  col- 
leagues at  both  the  University  of  Alberta  and 
the  California  Academy  of  Sciences  for  their  en- 
couragement and  assistance.  Special  thanks  are 
due  to  P.  H.  Arnaud,  Jr.,  G.  E.  Ball,  T.  L.  Er- 
win,  H.  Goulet,  G.  R.  Noonan,  and  D.  R.  White- 
head  for  their  continued  interest  in  the  project 
and  their  willingness  to  share  with  me  their  own 
observations  and  experiences. 

This  study  was  supported  by  the  National  Re- 
search Council  of  Canada  with  funds  from  Grant 
A- 1399  to  G.  E.  Ball.  Additional  financial  sup- 
port was  obtained  from  the  Boreal  Institute  for 
Northern  Studies  (University  of  Alberta), 
through  a  Grant  In  Aid  of  Research  (1973-74) 
for  fieldwork  in  Alaska,  and  from  the  California 
Academy  of  Sciences. 

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FIGURE  1.    Nebria  schwarzi  beverlianna,  new  subspecies:  adult  male,  dorsal  aspect  (approximately  6  times  actual  size). 
(Illustration  by  C.  L.  Mullinex.) 


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121 


HL 


PL 


EL 


FIGURES  1-11.  Fig.  1.  Components  of  "standardized  body  length"  measurement:  HL  =  head  length;  PL  =  pronotal  length; 
EL  =  elytral  length.  Figs.  2-8.  Right  antennal  scape,  dorsal  aspect.  2.  Nebria  gouleti  n.sp.  (Underwood,  Washington).  3.  Nebria 
gyllenhali  lassenensis  n.ssp.  (a.  Mount  Lassen,  California;  b.  Todd  Lake,  Oregon).  4.  Nebria  gyllenhali  lindrothi  n.ssp.  (a. 
Brooklyn  Lake,  Wyoming:  b.  Rio  Puerco,  New  Mexico).  5.  Nebria  zioni  oasis  n.ssp.  (Pine  Valley  Mountains,  Utah).  6.  Nebria 
spatulata  sierrae  n.ssp.  (Big  Horn  Lake,  California).  7.  Nebria  vandykei  wyeast  n.ssp.  (Mount  Hood,  Oregon).  8.  Nebria 
trifaria  utahensis  n.ssp.  (Henry  Mountains,  Utah).  Figs.  9-11.  Mentum,  ventral  aspect.  9.  General  form  and  chaetotaxy; 
M,  through  M4  are  designations  for  specific  pairs  of  setae.  10.  Nebria  carri  n.sp.  (Dollarhide  Summit,  Idaho).  11.  Nebria 
kincaidi  balli  n.ssp.  (Mount  Hood,  Oregon).  All  scale  lines  equal  1.0  mm. 


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am 


mis 


FIGURES  12-16.  Fig.  12.  Pronotum,  general  form  and  associated  terms  (a.  dorsal  aspect;  b.  left  lateral  aspect;  c.  cross-sec- 
tional aspect);  abbreviation  code:  aa  =  apical  angle;  am  =  apical  margination;  ati  =  anterior  transverse  impression;  ba  =  basal 
angle;  bf  =  basal  fovea;  bis  =  basolateral  seta;  Ib  =  lateral  bead  (=  lateral  margination);  le  =  lateral  explanation;  Ig  =  lateral 
groove;  mli  =  median  longitudinal  impression;  mis  =  midlateral  seta;  pep  =  proepipleuron;  pti  =  posterior  transverse  impres- 
sion. Figs.  13-16.  Pronotum,  dorsal  aspect.  13.  Nebria  gouleti  n.sp.  (Salmon  River,  Idaho).  14.  Nebria  lacustris  bellorum  n.ssp. 
(Great  Balsam  Mountains,  North  Carolina).  15.  Nebria  nivalis  gaspesiana  n.ssp.  (Mont  Albert,  Quebec).  16.  Nebria  acuta 
quileute  n.ssp.  (Olympic  Hot  Springs,  Washington).  Scale  line  equals  1.0  mm. 


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123 


FIGURES  17-24.  Pronotum,  dorsal  aspect.  17.  Nebria  sahlbergii  modoc  n.ssp.  (Warner  Mountains,  California).  18.  Nebria 
sahlbergii  triad  n.ssp.  (Trinity  Alps,  California).  19.  Nebria  ohliaua  rhuskae  n.ssp.  (Chuska  Mountains,  Arizona).  20.  Nebria 
darlingtoni  n.sp.  (Kyburz,  California).  21.  Nebria  gebleri  cascadensis  n.ssp.  (Glacier,  Washington).  22.  Nebria  gebleri  fra- 
gariae  n.ssp.  (Strawberry  Mountains,  Oregon).  23.  Nebria  gebleri  siskiyouensis  n.ssp.  (Trinity  Alps,  California).  24.  Nebria 
carri  n.sp.  (Dollarhide  Summit,  Idaho).  Scale  line  equals  1.0  mm. 


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FIGURES  25-29.  Figs.  25-28.  Pronotum,  dorsal  aspect.  25.  Nebria  meanyi  lamarckensis  n.ssp.  (Upper  Lamarck  Lake,  Cali- 
fornia). 26.  Nebria  meanyi  sylvatica  n.ssp.  (Olympic  Hot  Springs,  Washington).  27.  Nebria  schwarzi  beverlianna  n.ssp.  (Hoback 
River,  Wyoming).  28.  Nebria  trifaria  utahensis  n.ssp.  (Henry  Mountains,  Utah).  Fig.  29.  Prosternal  intercoxal  process,  Nebria 
lituyae  n.sp.  (Mount  Blunt,  Lituya  Bay,  Alaska).  All  scale  lines  equal  1.0  mm. 


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125 


35 


FIGURES  30-36.  Figs.  30-32.  Basal  region  of  left  elytron,  dorsal  aspect.  30.  Nehria  arkansana  edwardsi  n.ssp.  (a.  Rancheria, 
Yukon  Territory:  b.  Gorge  Creek,  Alberta).  31.  Nebria  kincaidi  balli  n.ssp.  (Mount  Rainier,  Washington).  32.  Nehria  meanyi 
lamarckensis  n.ssp.  (Upper  Lamarck  Lake,  California).  Figs.  33-35.  Left  hindwing.  33.  Full-sized  wing,  Nehria  arkansana 
edwardsi  n.ssp.  (Logan  Pass,  Montana).  34.  Wing  reduced  in  length,  Nebria  obliqua  chuskae  n.ssp.  (Chuska  Mountains, 
Arizona).  35.  Wing  reduced  in  length  and  width,  Nebria  arkansana  uinta  n.ssp.  (Logan  River,  Utah).  Fig.  36.  Right  hind  coxa, 
ventral  aspect,  Nebria  schwarzi  beverlianna  n.ssp.  (Hoback  River,  Wyoming).  All  scale  lines  equal  1.0  mm. 


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FIGURES  37-40.  Fig.  37.  Abdominal  venter,  form  and  associated  terms  (a.  ventral  aspect;  b.  left  lateral  aspect);  abbreviation 
code:  AS  =  anal  sternum;  HC  =  hind  coxa;  HF  =  hind  femur;  LP  =  lateral  pit;  MTE  =  metepisternum;  MTS  =  metaster- 
num;  PPMP  =  posterior  paramedial  puncture  and  associated  seta;  VS1  to  VS5  =  first  to  fifth  visible  abdominal  sterna.  Figs. 
38-40.  Median  lobe  of  male  genitalia.  38.  Nebria  acuta  quileute  n.ssp.  (Olympic  Hot  Springs,  Washington)  (a.  left  lateral 
aspect:  b.  ventral  aspect,  apical  region  only).  39.  Nebria  arkansana  edwardsi  n.ssp.  (Yoho  National  Park,  British  Columbia) 
(a.  left  lateral  aspect;  b.  dorsal  aspect,  apical  region  only;  c.  cross  section  at  middle  of  shaft).  40.  Nebria  arkansana  oowah 
n.ssp.  (La  Sal  Mountains,  Utah)  (left  lateral  aspect).  All  scale  lines  equal  1.0  mm. 


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127 


FIGURES  41-45.  Median  lobe  of  male  genitalia,  left  lateral  aspect.  41.  Nebria  arkansana  uinta  n.ssp.  (Logan  River,  Utah). 
42.  Nehriafragilis  teewinot  n.ssp.  (Mount  Teewinot,  Wyoming).  43.  Nehria  navajo  n.sp.  (19  miles  [ca.  31  km]  sw  of  Kayenta, 
Arizona).  44.  Nebria  gebleri  fragariae  n.ssp.  (Strawberry  Mountains,  Oregon).  45.  Nebria  trifaria  utahensis  n.ssp.  (Henry 
Mountains,  Utah).  Scale  line  equals  1.0  mm. 


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FIGURES  46-50.  Bursa  copulatrix  (a.  dorsal  aspect;  b.  left  lateral  aspect;  c.  bursal  sclerite  enlarged,  dorsal  aspect;  d.  bursal 
sclerite  enlarged,  left  lateral  aspect);  in  a.  or  b.,  stippled  area  denotes  shape  and  location  of  distinct  sclerite  or  moderately  to 
markedly  sclerotized,  but  less  clearly  defined,  area;  in  c.  or  d.,  stippled  area  denotes  weakly  sclerotized  membrane,  sclerite  is 
not  stippled.  46.  Nebria  acuta  quileute  n.ssp.  (Olympic  Hot  Springs,  Washington).  47.  Nebria  arkansana  edwardsi  n.ssp. 
(Amiskwi  Falls,  British  Columbia).  48.  Nebria  arkansana  oowah  n.ssp.  (La  Sal  Mountains,  Utah).  49.  Nebria  geblerifragariae 
n.ssp.  (Strawberry  Mountains,  Oregon).  50.  Nebria  trifaria  utahensis  n.ssp.  (Henry  Mountains,  Utah).  Scale  line  "x"  equals 
1.0  mm  and  applies  to  all  figures  "a"  and  "b."  Scale  line  "y"  equals  0.10  mm  and  applies  to  all  figures  "c"  and  "d." 


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129 


FIGURES  51-52.     Geographical  distribution  maps.   51.  Nebria  gouleti  n.sp.   52.  Nebria  lacustris  Casey  [N.  lacustris  lacus- 
tris  =  solid  circles;  N.  lacustris  bellorum  n.ssp.  =  solid  squares].  All  scale  lines  equal  500  km. 


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FIGURES  53-54.  Geographical  distribution  maps.  53.  Nebria  nivalis  (Paykull)  [N.  nivalis  nivalis  =  solid  triangles;  N.  nivalis 
gaspesiana  n.ssp.  =  solid  circles].  54.  Nebria  gyllenhali  (Schonherr)  [N.  gyllenhali  castanipes  (Kirby)  =  solid  circles;  N. 
gyllenhali  lassenensis  n.ssp.  =  solid  squares;  N.  gyllenhali  lindrothi  n.ssp.  =  solid  triangles].  All  scale  lines  equal  500  km. 


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131 


FIGURES  55-59.  Geographical  distribution  maps.  55.  Nebria  acuta  Lindroth  [N.  acuta  acuta  =  solid  circles;  N.  acuta 
quileute  n.ssp.  =  solid  triangles].  56.  Nebria  sahlbergii  Fischer  von  Waldheim  [Nebria  sahlbergii  sahlbergii  =  solid  circles; 
N.  sahlbergii  modoc  n.ssp.  =  solid  triangles;  N.  sahlbergii  triad  n.ssp.  =  solid  square].  57.  Nebria  lituyae  n.sp.  58.  Nebria 
arkansana  Casey  [N.  arkansana  arkansana  =  solid  triangles;  N.  arkansana  edwardsi  n.ssp.  =  solid  circles;  N.  arkansana 
oowah  n.ssp.  =  open  square;  N.  arkansana  uinta  =  open  triangles].  59.  Nebria  fragilis  Casey  [N.  fragilis  fragilis  =  solid 
circles;  N.  fragilis  teewinot  n.ssp.  =  solid  triangles].  All  scale  lines  equal  500  km. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  4 


FIGURES  60-65.  Geographical  distribution  maps.  60.  Nebria  zioni  Van  Dyke  [N.  zioni  zioni  =  solid  triangles;  N.  zioni  oasis 
n.ssp.  =  solid  circle].  61.  Nebria  obliqua  LeConte  [N.  obliqua  obliqua  -  solid  circles;  N.  obliqua  chuskae  n.ssp.  =  solid 
triangle].  62.  Nebria  darlintoni  n.sp.  63.  Nebria  navajo  n.sp.  64.  Nebria  gebleri  Dejean  [N.  gebleri  gebleri  -  solid  squares; 
N.  gebleri  cascadensis  n.ssp.  =  solid  triangles;  N.  gebleri  fragariae  n.ssp.  =  open  square;  N.  gebleri  rathvoni  LeConte  = 
solid  circles;  N.  gebleri  siskiyouensis  =  open  triangles].  65.  Nebria  carri  n.sp.  [solid  squares]  and  Nebria  kincaidi  Schwarz 
[N.  kincaidi  kincaidi  =  solid  circles;  N.  kincaidi  balli  n.ssp.  =  solid  triangles].  All  scale  lines  equal  500  km. 


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133 


FIGURES  66-70.  Geographical  distribution  maps.  66.  Nebria  spatulata  Van  Dyke  [N.  spatulata  spatulata  =  solid  circles; 
N.  spatulata  sierrae  n.ssp.  =  solid  triangles].  67.  Nebria  meanyi  Van  Dyke  [N.  meanyi  meanyi  =  solid  circles;  N.  meanyi 
lamarckensis  n.ssp.  =  solid  square;  N.  meanyi  sylvatica  n.ssp.  =  solid  triangles].  68.  Nebria  vandykel  Banninger  [N.  vandykei 
vandykei  =  solid  circles;  N.  vandykei  wyeast  n.ssp.  =  solid  triangles].  69.  Nebria  schwarzi  Van  Dyke  [N.  schwarzi  schwarzi  = 
solid  circles;  N.  schwarzi  beverlianna  n.ssp.  =  solid  triangle].  70.  Nebria  trifaria  LeConte  [N.  trifaria  trifaria  =  solid  circles; 
N.  trifaria  catenata  Casey  =  solid  triangles;  N.  trifaria  utahensis  n.ssp.  =  solid  square].  All  scale  lines  equal  500  km. 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  5,  pp.  135-179 


December  22,  1979 


EASTERN  PACIFIC  MACROURINE  GRENADIERS  WITH  SEVEN 
BRANCHIOSTEGAL  RAYS  (PISCES:  MACROURIDAE) 

By 

Tomio  Iwamoto 

Department  of  Ichthyology,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  94118 


ABSTRACT:  Seventeen  species  representing  eight  genera  of  macrourine  grenadiers  with  seven  branchiostegal 
rays  are  treated  as  part  of  the  eastern  Pacific  fauna.  Among  the  eight  genera,  Nezumia  is  the  most  diverse,  with 
10  species  represented.  Nezumia  ventralis  new  species,  is  described  from  two  Galapagos  specimens;  the  high 
pelvic  fin  ray  count  of  15  contrasts  this  species  with  all  other  eastern  Pacific  Nezumia.  Two  apparently  disjunct 
populations  of  \ .  loricata  are  given  subspecific  recognition:  subspecies  loricata  from  the  Galapagos  and  sub- 
species atomos  (new)  from  central  Chile.  Echinomacrurus,  Hymenocephalus,  Paracetonurus,  Ventrifossa,  Mal- 
acocephalus,  Mataeocephalus,  and  Mesobius  are  each  represented  by  only  one  species — the  first  four  genera 
listed  are  first  recorded  from  the  eastern  Pacific.  Echinomacrurus  occidentalis  (a  second  species  in  the  genus) 
is  newly  described  from  a  single  specimen  taken  off  Peru  in  4,334  m.  Macrurus  fragilis  Garman,  1899,  is 
tentatively  aligned  with  members  of  Paracetonurus.  Ventrifossa  is  recognized  as  consisting  of  three  subgenera: 
Ventrifossa,  Lucigadus,  and  Sokodara  (new).  Only  subgenus  Lucigadus  is  represented  in  the  eastern  Pacific. 


INTRODUCTION 

The  large  grenadier  subfamily  Macrourinae  is 
divisible  into  two  distinct  groups  based  on  bran- 
chiostegal ray  counts.  Those  macrourines  with 
six  branchiostegal  rays,  typified  by  Macrourus 
Bloch  and  Coryphaenoides  Gunnerus,  form  one 
group;  those  with  seven  branchiostegal  rays  and 
one  monotypic  genus  (Pseudonezumia  Okamu- 
ra)  with  seven  or  eight  form  a  second  group. 
This  second  group  may  be  further  divided  by 
species  differences  in  the  development  and  lo- 
cation of  the  periproct  region  (the  specialized 
area  surrounding  the  anal  and  urogenital  open- 
ings) and  the  development  of  ventral  light  or- 
gans. Hymenocephalus  is  the  most  distinctive 
member  of  this  second  group  and  forms  a  phy- 
logenetic  line  well  removed  from  the  others. 
Another  line  of  related  genera,  characterized  by 
the  members  having  a  broad  periproct  situated 


close  to  the  origin  of  the  anal  fin,  includes  such 
diverse  genera  as  Echinomacrurus  Roule,  Par- 
acetonurus Marshall,  Cetonurus  Giinther, 
Trachonurus  Giinther,  and  Sphagemacrurus 
Fowler.  These  genera  contrast  with  Nezumia 
Jordan,  Malacocephalus  Giinther,  Ventrifossa 
Gilbert  and  Hubbs,  Kumba  Marshall,  and  Pseu- 
donezumia Okamura,  members  of  which  have 
a  smaller  periproct  removed  by  some  distance 
from  the  origin  of  the  anal  fin  and  preceded  by 
an  anteriorly  extended  light  organ  of  various 
size.  Mataeocephalus  Berg  falls  somewhere  be- 
tween these  two  divisions  in  that  members  of 
one  species  (M.  adjustus)  have  a  periproct  that 
is  well  removed  from  the  anal  fin  origin,  and 
they  also  have  a  distinct  dermal  window  of  the 
light  organ  extended  forward  of  the  periproct, 
while  members  of  the  other  species  apparently 
have  a  periproct  situated  close  before  the  anal 


[135] 


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fin,  and  their  light  organ  lacks  a  distinct  dermal 
window. 

The  16  species  here  considered  constitute 
about  40  percent  of  the  total  (approximately  40 
spp.)  macrourid  fauna  of  the  eastern  Pacific.  The 
genus  Coelorinchus  from  this  region  (6  spp.) 
was  reviewed  in  a  previous  paper  (Iwamoto 
1979).  A  review  of  the  large  complex  of  ma- 
crourines  with  six  branchiostegal  rays  and  the 
trachyrincines  (2  spp.)  from  the  eastern  Pacific 
is  currently  underway.  A  summary  of  past  stud- 
ies of  the  Macrouridae  from  eastern  Pacific 
waters  is  given  in  Iwamoto  and  Stein  (1974).  In 
addition,  the  recent  works  of  Hubbs  and  Iwa- 
moto (1977),  Chirichigno  and  Iwamoto  (1976), 
Shcherbachev  et  al.  (1979),  Ojeda  and  Camus 
(1977),  and  Parin  et  al.  (1973,  1976)  should  be 
consulted. 

METHODS 

Methods  for  making  counts  and  measure- 
ments generally  follow  procedures  described  by 
Hubbs  and  Lagler  (1958)  and  modified  for  ma- 
crourids  by  Iwamoto  (1970,  1978).  Six  or  seven 
branchiostegal  rays  are  found  in  all  but  one 
species  of  macrourid.  The  exception  is  Pseu- 
donezumia  Okamura,  197 la,  which  has  seven  or 
eight  (personal  communications,  Osamu  Oka- 
mura, September  1978).  Four  of  the  rays  are  at- 
tached laterally  on  the  epihyal  and  ceratohyal; 
the  remaining  two  or  three  (or  probably  four  in 
Pseudonezumia;  condition  not  known)  are  at- 
tached anteriorly  and  medially  on  the  cerato- 
hyal— thus  only  the  latter  group  of  rays  need  be 
counted  to  determine  the  total  number.  How- 
ever, these  anteriormost  rays  (i.e.,  those  closest 
to  the  isthmus)  are  generally  small  and  slender, 
making  them  difficult  to  see  without  probing  or 
dissection. 

Synonymies  are  limited  to  primary  synonyms 
and  other  combinations.  Materials  for  this  study 
are  based  primarily  on  collections  made  by  the 
ANTON  BRUUN  in  1966  and  the  TE  VEGA  in  1968; 
most  of  these  are  deposited  in  the  ichthyological 
collection  of  the  California  Academy  of  Sciences 
(CAS).  Other  sources  have  been  extensively 
used,  and  abbreviations  for  the  depositories  are 
as  follows:  AMNH,  American  Museum  of  Nat- 
ural History,  New  York;  BMNH,  British  Mu- 
seum of  Natural  History,  London;  CAS-SU, 
Natural  History  Museum,  Stanford  University, 
now  housed  at  CAS;  FAKU,  Faculty  of  Agri- 
culture, Kyoto  University,  Maizuru;  FSFRL, 


ventral  striae 


ventral   striae 


smooth 


fragile  bones 


FIGURE  1.  Diagrammatic  ventral  (a)  and  lateral  (b)  views 
of  a  hypothetical  representative  of  the  genus  Hymenocepha- 
lus,  showing  diagnostic  features  of  the  genus. 


Far  Seas  Fisheries  Research  Laboratory,  Shi- 
mizu;  IMARPE,  Institute  del  Mar,  Callao; 
LACM,  Natural  History  Museum  of  Los  An- 
geles County,  Los  Angeles;  MCZ,  Museum  of 
Comparative  Zoology,  Harvard  University, 
Cambridge;  SIO,  Scripps  Institution  of  Ocean- 
ography, La  Jolla;  UMML,  School  of  Marine 
and  Atmospheric  Science,  University  of  Miami, 
Miami;  USNM,  National  Museum  of  Natural 
History,  Washington,  D.C. 


KEY  TO  THE  ADULTS  OF  EASTERN  PACIFIC 

MACROURINE  SPECIES  WITH  SEVEN 

BRANCHIOSTEGAL  RAYS 

la.  Ventral  striae  (fine  black  transverse 
lines,  most  readily  visible  under  magni- 
fication) present  over  gular  membranes, 
isthmus,  and  parts  of  chest  and  abdomen 
(Fig.  1).  Anus  situated  immediately  be- 
fore anal  fin  and  preceded  by  a  small. 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


137 


smooth 


scales  on 
branchiostegal   rays 

FIGURE  2.  Malacocephalus  laevis.  (a)  Lateral  view  show- 
ing smooth  leading  edge  of  second  spinous  dorsal  ray  and 
enlarged  teeth  of  lower  jaw.  (b)  Ventral  view  showing  scales 
on  branchiostegal  rays  and  location  of  anus  and  light  organ 
structures. 


raised  lenslike  structure;  this  structure 
connected  by  a  thin  (often  obscure) 
black  median  line  to  similar  structure  on 
chest.  Head  bones  extremely  fragile, 
some  almost  membranous.  Gill-rakers 
numerous,  more  than  20  (total  count)  on 
first  arch.  Second  spinous  dorsal  ray 

smooth Hymenocephalus  sp.  (p.  140) 

Ib.  No  ventral  striae.  Anus  removed  by  a 
short  to  moderate  distance  from  anal  fin; 
lenslike  structure,  if  present,  far  re- 
moved from  anal  fin  origin  and  usually 
located  in  a  shallow  fossa.  Head  bones 
relatively  strong.  Gill-rakers  fewer  than 
20  (total)  on  first  arch.  Second  spinous 
dorsal  ray  smooth  or  serrated  2 

2a.  Spinous  second  ray  of  first  dorsal  fin 
with  smooth  leading  edge  (Fig.  2a) 3 

2b.  Spinous  second  ray  of  first  dorsal  fin 
with  serrated  leading  edge  5 

3a.  Chin  barbel  absent  „ 


a 


broad  naked   margin 


dermal   window 


Mesobius  berryi  Hubbs 
and  Iwamoto  (p.  141) 


FIGURE  3.  Diagrammatic  ventral  views  of  abdomen  of 
grenadiers  showing  periproct  location  (a)  abutting  or  close  to 
(separated  by  a  few  scale  rows  in  some  individuals)  origin  of 
anal  fin,  and  (b)  periproct  far  removed  from  origin  of  anal  fin. 


3b.  Chin  barbel  present,  well  developed  __._   4 

4a.  Teeth  large,  in  a  single  row  on  lower  jaw 
(Fig.  2a);  scales  on  branchiostegal  mem- 
brane (Fig.  2b)  

__  Malacocephalus  laevis  (Lowe)  (p.  149) 
4b.  Teeth  small,  in  a  narrow  band  on  lower 
jaw;  no  scales  on  branchiostegal  mem- 
branes   

Nezumia  liolepis  (Gilbert)  (p.  157) 

5a.  Periproct  close  to  or  abutting  (Fig.  3a) 
origin  of  anal  fin;  anus  usually  closer  to 
origin  of  anal  fin  than  to  insertion  of  pel- 
vic fins  6 

5b.  Periproct  far  removed  from  origin  of  anal 
fin  (Fig.  3b);  anus  usually  closer  to  in- 


138 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  5 


FIGURE  4.  Ventrolateral  views  of  (a)  Ventrifossa  nigro- 
maculata  and  (b)  Nezumia  latirostrata  comparing  relative 
lengths  of  first  dorsal  fin,  anterior  extent  of  gill  openings  (ar- 
rows), and  relative  positions  of  pelvic  and  anal  fins. 


sertion  of  pelvic  fins  than  to  origin  of 
anal  fin 8 

6a.  Head  massive,  globose;  snout  broadly 
rounded.  Scales  on  body  distinctly  non- 
imbricate  (not  overlapping),  the  exposed 
field  of  each  scale  separated  by  a  fleshy 
border  from  the  exposed  fields  of  adja- 
cent scale 

Echinomacrurus  occidentalis  n.  sp.  (p.  143) 

6b.  Head  relatively  slender,  angular;  snout 
pointed.  Body  scales  distinctly  imbri- 
cate; exposed  fields  of  adjacent  scales 
not  distinctly  separated  from  each  other 
by  a  fleshy  border 7 

7a.  Head  bones  stout;  head  ridges  strongly 
armed  with  coarse  scutelike  scales. 
Mouth  small,  notably  inferior,  and  dis- 
tinctly U-shaped.  Scales  densely  cov- 
ered with  spinules  


Mataeocephalus  tenuicauda 

(Carman)  (p.  145) 

7b.  Head  bones  fragile;  head  ridges  naked  or 
with  unmodified  scales.  Mouth  moderate 
in  size,  not  notably  inferior,  and  more 
normally  shaped.  Scales  with  few  or  no 

spinules  on  exposed  field 

Paracetonurus  fragilis 

(Carman)  (p.  147) 

8a.  Height  of  first  dorsal  fin  much  greater 
than  length  of  head.  Opercular  opening 
extends  far  forward  to  vertical  through 

hind  edge  of  maxillae  (Fig.  4a) 

„  Ventrifossa  (Lucigadus)  nigromaculata 
(McCulloch)  (p.  153) 

8b.  Height  of  first  dorsal  fin  about  equal  to 
or  less  than  length  of  head.  Anteriormost 
extent  of  opercular  opening  much  pos- 
teriad  of  vertical  through  hind  edge  of 
maxillae  (Fig.  4b) 9 

9a.  Second  ray  of  first  dorsal  fin  with  0  to  4 
weak  denticles  on  leading  edge  (Fig.  5a). 
Scales  thin,  highly  deciduous  with  few 
or  no  spinules  on  exposed  fields  of  body 

scales  

_r Nezumia  liolepis  (Gilbert)  (p.  157) 

9b.  Second  ray  of  first  dorsal  fin  with  nu- 
merous prominent  denticulations  on 
leading  edge  (Fig.  5b).  Scales  relatively 
adherent,  with  spinules  densely  covering 
exposed  fields 10 

lOa.  Species  bathypelagic.  Gill  filaments 
short;  length  about  half  diameter  of  eye 
lens.  Outer  pelvic  ray  prolonged,  70- 
160%  HL.  Body  scales  small,  bearing  1- 
15  slender,  erect  spinules  that  render 
body  surface  velvety;  11-14  scale  rows 
below  origin  of  second  dorsal  fin.  Color 

black  to  brownish  black  

Nezumia  parini  Hubbs  and 

Iwamoto  (p.  176) 

lOb.  Species  benthopelagic.  Gill  filaments 
moderate  to  long;  length  about  equal  to 
or  greater  than  diameter  of  eye  lens. 
Outer  pelvic  ray  usually  less  than  70% 
HL.  Body  scales  small  to  moderate;  spi- 
nules on  scales  few  to  numerous,  mod- 
erately to  greatly  inclined,  conical,  lan- 
ceolate, or  shield  shaped,  usually  rough 
in  texture;  fewer  than  1 1  scale  rows  be- 
low origin  of  second  dorsal  fin.  Color 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


139 


a 


FIGURE  5.  Comparison  of  denticulations  on  leading  edge 
of  second  spinous  dorsal  rays  of  (a)  Nezumia  liolepis  and  (b) 
Nezumia  loricata. 


various  shades  of  blue,  violet,  brown,  or 
black.,.  11 

1  la.  Mandibular  rami  usually  completely  na- 
ked; pores  of  lateralis  system  large  and 
prominent  on  mandibular  rami  and  along 
ventral  border  of  suborbital  region  (Fig. 
6a).  Mouth  relatively  large,  length  upper 
jaw  usually  30-36%  HL.  Outer  gill-slit 
relatively  wide,  17-22%  HL  ..  12 

lib.  Most  of  mandibular  rami  scaled,  al- 
though anterior  end  naked  in  some; 
pores  of  cephalic  lateralis  system  small, 
relatively  inconspicuous  (Fig.  6b).  Mouth 
relatively  small,  length  upper  jaws  23- 
32%  HL.  Length  outer  gill-slit  12-16% 
HL.  13 


12a.  Patches  of  small  scales  on  base  of  low- 
ermost branchiostegal  rays  (Fig.  6a). 
Spinules  on  body  scales  relatively 
broad,  lanceolate,  arranged  in  a  some- 
what quincunx  pattern,  rows  not  dis- 
cretely parallel  or  slightly  convergent 
and  ridgelike.  Chin  barbel  long,  20-25% 
HL.  Rays  of  pelvic  fin  10-11.  Orbit  di- 
ameter 25-29%  HL  „ 
__  Nezumia  stelgidolepis  (Gilbert)  (p.  161) 

12b.  Few  isolated  scales  or  no  scales  on  bran- 
chiostegal rays.  Spinules  on  body  scales 
narrow,  usually  conical,  arranged  in  dis- 
crete parallel  or  slightly  convergent 
rows.  Length  of  chin  barbel  10-16%  HL. 
Rays  of  pelvic  fin  1 1-12.  Orbit  diameter 
30-34%  HL  . 
— _  Nezumia  pulchella  (Pequeno)  (p.  159) 

13a.  Gill-rakers  on  inner  side  of  first  (outer- 
most) arch  9  or  fewer  (total  count),  5-6 

(rarely  7)  on  lower  limb  

__  Nezumia  convergens  (Garman)  (p.  171) 

13b.  Gill-rakers  on  inner  side  of  first  arch  9 
or  more  (total),  8  or  9  on  lower  limb  ____ 

14 

14a.   Pelvic  fin  rays  9.  First  dorsal  with  prom- 
inent black  blotch  distally,  pale  basally. 
Interorbital  region  narrow,  width   17- 
19%  HL... 
Nezumia  orbitalis  (Garman)  (p.  167) 

14b.  Pelvic  fin  rays  9-12.  First  dorsal  fin  usu- 
ally uniformly  blackish.  Interorbital 
width  moderate,  17-26%  (usually  more 
than  20%)  HL-  15 


a  b 

FIGURE  6.     Ventrolateral  views  of  (a)  Nezumia  stelgidolepis  and  (b)  Nezumia  pudens  comparing  pore  development  (slightly 
exaggerated)  and  squamation  of  head  in  the  two  species. 


140 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  5 


15a.  Snout  relatively  blunt,  high.  Suborbital 
region  smoothly  and  almost  completely 
scaled;  scales  along  ridges  not  especially 
stout  and  coarse  (Fig.  6b).  Barbel  long, 

19-25%  HL  Neiumia  pudens 

Gilbert  and  Thompson  (p.  163) 

15b.  Snout  pointed,  low.  Suborbital  region 
often  with  ventral  surfaces  partially  na- 
ked; scales  along  ridges  stout,  coarse. 
Barbel  shorter,  less  than  17%  HL  ____  16 

16a.  Pelvic  fin  rays  9-10.  First  dorsal  fin  rays 
11,8-11  (usually  9-10)  Length  barbel  5- 

10%  HL 

..  Nezumia  latirostrata  (Garman)  (p.  168) 

16b.  Pelvic  fin  rays  10-12.  First  dorsal  fin 
rays  11,9-12  (usually  10-12).  Length  bar- 
bel 10-17%  HL 

Nezumia  loricata  (Garman)  (p.  174) 


Hymenocephalus  Giglioli 

Hymenocephalus  GIGLIOLI,  1882:199  (type-species  Hymeno- 
cephalus italicus  Giglioli,  1884,  by  monotypy). 

Mystaconurus  GUNTHER,  1887: 124  (as  subgenus  of  Macrurus) 
(type-species  Hymenocephalus  italicus  Giglioli,  1884,  by 
subsequent  designation  of  Jordan  and  Evermann  1898:2580). 

Hymenogadus  GILBERT  AND  HUBBS,  1920:521  (as  subgenus) 
(type-species  Hymenocephalus  gracilis  Gilbert  and  Hubbs, 
1920,  by  original  designation).  — Okamura  1970a:58  (rec- 
ognized as  genus). 

DIAGNOSIS. — Macrourine  grenadiers  with  anus 
immediately  before  anal  fin.  Striae,  consisting  of 
fine  parallel  black  lines  over  silvery  pigment,  on 
most  ventral  areas  of  body.  Two  lenslike  dermal 
windows  of  light  organ  on  midventral  line;  one 
immediately  before  anus  and  one  on  chest  be- 
fore pelvic  fin  bases;  dermal  window  connected 
by  black  ridge  along  midventral  wall  of  abdom- 
inal cavity.  Head  large,  cavernous;  bones  thin, 
head  covering  membranous.  Mouth  large,  sub- 
terminal.  Gill  openings  and  outer  gill-slit  rela- 
tively wide.  Gill-rakers  numerous,  more  than  15 
on  lower  limb  of  outer  arch  except  in  slender, 
cylindrical-bodied  species  (subgenera  Hymeno- 
gadus and  Spicomacrurus).  Second  dorsal  spine 
smooth  or  weakly  denticulate  (in  Hymenoga- 
dus). Small  species,  usually  less  than  200  mm 
TL.  (Adapted  after  Iwamoto  1970:374-375.) 

REMARKS. — Okamura  (1970a)  elevated  the 
subgenus  Hymenogadus  to  full  generic  status, 
including  in  it  H.  gracilis  Gilbert  and  Hubbs, 
1920,  H.  tennis  Gilbert  and  Hubbs,  1917,  and 
H.  kuronumai  Kamohara,  1938.  The  three 


species  are  obviously  closely  related  and  form 
a  distinct  group  readily  distinguished  from  most 
other  Hymenocephalus  (sensu  lato),  but  it  is  not 
the  differences  between  the  species  groups  in 
Hymenocephalus  but  the  similarities  among 
them  that  are  most  striking.  Used  in  its  widest 
sense,  the  genus  Hymenocephalus  encompasses 
a  group  of  species  that  is  notably  distinct  and 
distantly  removed  from  other  groups  of  ma- 
crourine  grenadiers.  By  recognizing  both  Hy- 
menocephalus and  Hymenogadus,  differences 
are  emphasized  and  the  obvious  close  relation- 
ship of  the  two  groups  is  masked.  This  is  re- 
grettable and  entails  a  needless  proliferation  of 
names.  Recognition  of  Hymenogadus  (with  gra- 
cilis and  tenuis)  and  Spicomacrurus  (with  ku- 
ronumai) as  subgenera  within  Hymenocephalus 
is  to  me  a  more  practicable  and  meaningful  treat- 
ment of  the  species  groups  involved. 

Hymenocephalus  sp. 

DIAGNOSIS. — Refer  to  generic  diagnosis. 

COUNTS  AND  MEASUREMENTS. —  ID.  11,8; 
gill-rakers  on  outer  arch  about  20.  The  following 
in  millimeters:  estimated  HL  25;  estimated  TL 
170;  horizontal  orbit  diameter  10.3;  length  upper 
jaw  15.0;  width  suborbital  3.8;  orbit  to  angle  of 
preopercle  12.3;  length  barbel  3.7. 

REMARKS. — The  single  specimen  of  this  genus 
from  eastern  Pacific  waters  was  in  such  poor 
condition  when  examined  in  June  1975,  that  an 
adequate  description  could  not  be  prepared.  The 
head  was  damaged  and  had  become  separated 
from  the  trunk,  and  the  paired  fins  were  gone; 
but  there  was  no  question  as  to  its  genus  because 
of  the  presence  of  striae  on  the  gular  membrane 
and  along  the  abdomen;  the  presence  of  two 
lenslike  light  organs  on  the  chest  and  before  the 
anus;  the  number,  shape,  and  coloration  of  the 
gill-rakers;  and  the  shape  of  the  preopercle 
ridge — these  combined  features  uniquely  char- 
acterize the  genus  Hymenocephalus. 

This  specimen  represents  the  first  record  of 
the  genus  from  eastern  Pacific  waters.  The  genus 
is  common  throughout  the  warm-water  regions 
of  the  Atlantic,  Indian,  and  most  of  the  central 
and  western  Pacific  oceans.  That  only  a  single 
specimen  has  been  collected — and  that  a  fair- 
sized  adult  taken  in  a  midwater  haul — suggests 
that  the  species  is  not  a  regular  inhabitant  of 
eastern  Pacific  waters.  The  specimen  may  rep- 
resent a  stray  or  an  expatriate,  possibly  from 
populations  far  to  the  westward. 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


141 


MATERIAL  EXAMINED. — USNM  149049  (1  specimen,  est. 
170  mm  TL);  Peru,  off  Aguja  Pt.,  5°57'30"S,  81°50'W;  vertical 
haul,  732-0  m  over  bottom  depth  of  4,023  m;  ALBATROSS  sta. 
4655,  12  Nov.  1904. 

Mesobius  Hubbs  and  Iwamoto 

Mesobius  HUBBS  AND  IWAMOTO,  1977:235  (type-species  Me- 
sobius berryi  Hubbs  and  Iwamoto,  1977,  by  original  desig- 
nation). 

DIAGNOSIS. — Bathypelagic.  Periproct  region 
broad,  somewhat  raised,  close  to  anal  fin  origin 
(removed  from  origin  by  2-4  scale  rows  in  some 
individuals);  anus  centrically  located  within 
periproct.  Large  light  organ  abutting  rectum,  but 
no  anterior  extensions  between  pelvic  fin  bases 
or  on  chest.  Abdomen  short,  distance  isthmus 
to  anal  fin  origin  1.0-1.5  of  orbit  diameter  in 
adults.  Chin  barbel  absent.  Dentition  in  both 
jaws  consist  of  narrow  bands  of  small  teeth. 
Scales  of  head  elongate,  each  bearing  1-3  rows 
of  spinules  that  form  low,  sharp  ridges  with  2- 
9  spinules  per  row.  Head  and  body  laterally 
compressed.  Swim  bladder  greatly  reduced, 
bearing  2  retia  and  2  gas  glands.  Postlarvae  and 
prejuveniles  pass  through  a  polka-dotted  ("phal- 
acromacrurus")  stage;  adults  mostly  black. 
(Adapted  from  original  description.) 

REMARKS. — Since  publication  of  the  genus 
description.  Dr.  Carl  L.  Hubbs  and  I  have  re- 
ceived information  concerning  additional  speci- 
mens of  the  genus.  Dr.  Nikolai  V.  Parin  has 
written  (to  Hubbs,  25  Nov.  1977)  that  represen- 
tatives of  the  genus  have  been  collected  by  Rus- 
sian vessels  in  the  Atlantic  and  Indian  oceans 
between  latitudes  31°  and  37°S.  These  specimens 
are  presumably  those  recently  reported  by 
Shcherbachev  et  al.  (1979)  who  recorded  M. 
berryi  from  the  Indian  Ocean,  and  M.  antipo- 
dum  from  the  Indian  Ocean  and  the  Atlantic 
Ocean  off  the  southwestern  tip  of  Africa.  Unlike 
the  type-specimens  of  both  species,  the  Russian 
specimens  were  captured  in  bottom  trawls.  The 
largest  M.  antipodum  recorded  by  Shcherba- 
chev et  al.  was  more  than  661  mm  in  total  length 
and  136  mm  in  head  length,  and  their  six  spec- 
imens ranged  104-136  mm  HL  and  465+  to  661  + 
mm  TL.  Their  five  specimens  of  M.  berryi 
ranged  64.5-75.5  mm  HL  and  330+  to  412+  mm 
TL.  The  M.  antipodum  specimens  are  consid- 
erably larger  than  any  of  the  M.  antipodum  and 
M.  berryi  Dr.  Hubbs  and  I  had  examined  (the 
largest  M.  berryi  we  had  was  392  mm  in  total 
length  and  70  mm  in  head  length;  the  holotype 


of  M.  antipodum  was  390  mm  long,  with  an  in- 
complete tail,  and  75  mm  in  head  length).  Dr. 
Christine  Karrer  has  also  informed  us  (personal 
communication,  Oct.  1977)  of  having  examined 
specimens  of  Mesobius  in  the  Institut  fur  See- 
fischerei  (ISH),  Hamburg,  that  were  captured  in 
the  Atlantic  by  the  WALTHER  HERWIG.  Four  of 
these  (ISH  1816/68;  88-107  mm  HL)  were  bor- 
rowed through  the  kindness  of  Dr.  M.  Stehman 
and  were  identified  as  M.  antipodum.  Takao 
Arai  ( 1979)  recently  reported  the  collection  of  12 
specimens  of  M.  antipodum  off  New  Zealand, 
the  largest  of  which  was  128  mm  in  head  length. 
It  thus  appears  that  M.  antipodum  attains  a 
much  larger  size  than  M.  berryi,  and  that  spec- 
ulation (Hubbs  and  Iwamoto  1977:246)  as  to  the 
probable  distribution  of  M.  antipodum  extend- 
ing widely  across  the  Southern  Hemisphere  is 
supported. 

Mesobius  berryi  Hubbs  and  Iwamoto 

(Figure  la) 

Mesobius  berryi  HUBBS  AND  IWAMOTO,  1977:236-244,  figs. 
1-8,  10A  (original  description;  holotype  and  18  paratypes 
from  eastern  North  Pacific). 

DIAGNOSIS. — A  species  of  Mesobius  with  12- 
17  pyloric  caeca;  mesial  gill-rakers  on  first  arch 
10-13  (x  =  11.8),  on  second  arch  10-12  (x  = 
11.0).  Length  posterior  nostril  less  than  4  into 
least  suborbital  width.  Differentiated  squama- 
tion  of  posttemporal  region  not  extending  pos- 
teriad  of  vertical  through  origin  of  pectoral  fin. 
Outer  margin  of  gill  cover  not  notably  incised  at 
subopercle. 

REMARKS. — General  features  of  the  fish  can 
be  seen  in  Figure  7a.  This  species  has  been  ad- 
equately described  in  the  original  description, 
and  in  the  recent  reports  by  Arai  (1979)  and 
Shcherbachev  et  al.  (1979).  Mesobius  berryi  and 
Nezumia  parini  are  the  only  eastern  Pacific 
species  of  macrourid  normally  living  bathype- 
lagically  as  adults  (adults  of  other  species  make 
excursions  into  bathypelagic  depths,  but  their 
normal  habitat  is  near  bottom,  i.e.,  benthope- 
lagic;  see  Marshall  and  Merrett  (1977)  and  Mer- 
rett  (1978)  for  recent  discussions  on  this  sub- 
ject). The  unique  squamation  of  the  head  of 
Mesobius  berryi  is  unlike  that  of  any  other 
species  encountered  in  the  eastern  Pacific  and 
makes  adults  of  the  species  easily  recognizable. 
The  specific  differences  originally  reported  be- 
tween M.  berryi  from  the  North  Pacific  and  M. 
antipodum  from  the  South  Pacific  have  generally 


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(I 


FIGURE  7.     (a)  Mesobius  berryi  Hubbs  and  Iwamoto.  (Adapted  from  fig.  1  in  Hubbs  and  Iwamoto  1977.)  (b)  Echinomacrurus 
occidentalis  Iwamoto,  new  species.  Reconstruction  of  holotype,  USNM  135612,  collected  by  the  ALBATROSS  off  Peru  in  4,334  m. 


been  confirmed  by  Arai  (1979)  upon  his  exami- 
nation of  12  specimens  of  M.  antipodum  cap- 
tured off  the  New  Zealand  shelf  and  by  Shcher- 
bachev  et  al.  (1979)  from  examination  of 
specimens  from  the  Indian  Ocean  and  the  south- 
western Atlantic.  Meristic  features  show  the 
best  differences  (see  Table  1),  with  the  pyloric 
caeca  count  showing  no  overlap  (12-17  in  eight 
specimens  of  M.  berryi,  30-43  in  more  than  six 
specimens  of  M.  antipodum)  (data  from  Hubbs 
and  Iwamoto  (1977),  Shcherbachev  et  al.  (1979), 
and  Arai  (1979));  the  last  author  gave  only  the 
range  of  counts  in  his  12  specimens  of  M.  an- 
tipodum). 

Echinomacrurus  Roule 

Echinomacrurus  ROULE,  1916:22  (type-species  Echinomacru- 
rus mollis  Roule,  1916,  by  monotypy). 

DIAGNOSIS. — Macrourine  grenadiers  with  anus 
located  within  a  broad  naked  area  that  lies  im- 
mediately adjacent  to  anal  fin  origin;  head  mas- 
sive, swollen  by  the  expansive  cephalic  lateral- 


line  canals;  second  spinous  ray  of  first  dorsal  fin 
serrated  along  leading  edge;  scales  nonimbri- 
cate,  with  slender,  erect  spinules;  scales  along 
dorsal  fins  not  especially  enlarged  (as  in  Ceto- 
nurus);  no  scales  on  gular  and  branchiostegal 
membranes;  swim  bladder  reduced  or  absent. 
(See  Marshall  (1973:599)  for  additional  charac- 
ters.) 

REMARKS. — The  genus  Echinomacrurus  has 
hitherto  been  known  only  from  the  eastern 
North  Atlantic  and  the  western  Indian  Ocean 
(Marshall  1973).  The  eastern  Pacific  specimen 
here  reported  is  the  first  Pacific  and,  at  4,334  m, 
the  shallowest  record  of  the  genus.  Other  spec- 
imens of  Echinomacrurus  have  been  taken  at 
depths  from  5,000  to  5,413  m,  and  except  for 
one  Indian  Ocean  specimen,  all  were  taken  in 
nets  fished  at  the  bottom — Marshall  (1973)  thus 
considers  E.  mollis  bathypelagic  to  benthope- 
lagic  in  habit. 

The  genus  is  closely  related  to  Cetonurus 
Vaillant,  1888;  the  two  agree  in  most  diagnostic 
characters  given  above  but  not  in  squamation 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


143 


TABLE  1.  COMPARISON  OF  SELECTED  COUNTS  IN  MESO- 
BWS  BERRYI  AND  M.  ANTlPODUM.  Data  from  Hubbs  and 
Iwamoto  (1977),  Aral  (1979),  and  Shcherbachev  et  al.  (1979). 

Total  gill-rakers  on  first  arch 


range 

X 

S.D. 

n 

M  .  berrvi 

10-13 

11.7 

0.82 

19 

M.  antipodum 

12-15 

13.9 

0.85 

31 

Total  gill-rakers  on  second  arch 


range 

X 

S.D. 

n 

M.  berryi 

10-12 

11.2 

0.62 

20 

M.  antipodum 

12-16 

13.9 

0.89 

31 

Pectoral  fin  rays 

range 

X 

S.D. 

n 

M  .  berrvi 

12-14 

12.9 

0.63 

33 

M.  antipodum 

13-16 

14.5 

0.68 

31 

Pelvic 

fin  rays 

range 

X 

S.D. 

n 

M.  berryi 

7-9 

7.7 

0.57 

35 

M.  antipodum 

6-7 

6.9 

0.35 

30 

and  swim-bladder  characters.  Cetonurus  has 
imbricate  scales  over  most  of  the  body  and  no- 
tably enlarged  scales  along  the  base  of  the  sec- 
ond dorsal  fin;  it  also  has  scales  over  the  lower 
branchiostegal  rays  and  on  the  gular  membrane, 
and  a  well-developed  swim  bladder. 

Echinomacrurus  occidentalis  new  species 
(Figure  7b) 

DIAGNOSIS. — An  Echinomacrurus  with  12 
pelvic  fin  rays.  Orbits  about  20  percent  HL. 
Eleven  gill-rakers  on  mesial  side  of  second  arch. 
Interspace  between  first  and  second  dorsal  fins 
31  percent  HL. 

COUNTS.— ID.  ca.  11,9;  IP.  il8/i!8;  2P.  ca.  127 
12.  Gill-rakers  on  first  arch  1  +  9;  on  second 
arch  2  +  9.  Pyloric  caeca  6. 

MEASUREMENTS  (all  in  mm;  measurements 
preceded  by  ca.  are  estimates). — Total  length 
ca.  400;  head  length  ca.  80;  snout  length  ca.  27; 
preoral  length  ca.  23;  horizontal  orbit  diameter 
16;  least  postorbital  length  39;  orbit  to  angle  of 
preopercle  36;  suborbital  width  10;  length  upper 
jaw  22.5;  length  barbel  7;  length  outer  gill-slit  9; 
preanal  length  ca.  110;  length  snout  to  anus  ca. 


102;  outer  pelvic  to  anal  29;  isthmus  to  anal  ca. 
54;  greatest  body  depth  ca.  62;  depth  over  anal 
origin  ca.  53;  1D.-2D.  interspace  31. 

DESCRIPTION. — Head  deep,  massive,  presum- 
ably swollen  in  appearance  when  fresh;  snout 
high,  broad;  nostrils  high,  located  at  about  level 
of  dorsal  margin  of  orbits;  interorbital  space 
broad,  convex.  Mouth  rather  small,  subinferior, 
but  upper  jaw  extends  near  to  vertical  through 
hind  edge  of  orbit;  ascending  limb  of  premaxilla 
high  (height  about  0.9  of  ramus  length),  inclined 
forward.  Interopercle  mostly  covered  by,  and 
closely  adhered  to,  preopercle.  A  distinct  notch 
in  outline  of  gill  cover  formed  by  posteroventral 
border  of  subopercle. 

Scales  small,  nonimbricate,  almost  in  mosaic 
pattern  on  parts  of  head,  more  widely  spaced  on 
body.  Thin,  erect  spinules  cover  most  scales, 
giving  shagreenlike  feel  to  skin.  Scales  every- 
where cover  exposed  surfaces  of  head  and  body 
except  on  fins,  lips,  parts  of  subopercle,  wide 
margin  surrounding  anus,  small  areas  behind 
pectoral  and  dorsal  fins,  and  gill  membranes. 
Shoulder  girdle  beneath  gill  cover  naked;  border 
between  naked  and  scaled  areas  of  shoulder 
sharply  demarcated.  Periproct  large,  raised,  im- 
mediately adjacent  to  anal  fin  and  occupying  al- 
most half  distance  between  insertion  of  pelvic 
fins  and  origin  of  anal  fin;  anus  slightly  protrud- 
ing. Premaxillary  teeth  small,  conical,  recurved, 
in  a  narrow,  tapered  band  of  about  four  rows 
deep  anteriorly,  tapering  to  a  single  row  poste- 
riorly; band  extends  about  three-fourths  length 
of  premaxillary  ramus;  outer  series  of  teeth 
slightly  enlarged.  Mandibular  dentition  about 
the  same  as  that  of  premaxillary,  but  without 
enlarged  outer  teeth. 

Gill-rakers  short,  spiny,  tubercular.  Outer  gill- 
slit  relatively  long;  about  8  rudimentary  rakers 
on  outer  side  of  first  arch,  1  +  9  on  inner  side, 
these  armed  with  slender  conical  spines.  Pyloric 
caeca  6,  large,  thick.  Specimen  a  male  with  2 
well-developed  testes. 

Linings  of  gill,  buccal,  and  peritoneal  cavities 
black.  Stomach  black. 

REMARKS. — It  is  with  some  reluctance  that  I 
describe  this  single  eastern  Pacific  specimen  of 
Echinomacrurus  as  a  new  species,  because  of 
its  extremely  poor  condition.  It  is  unlikely,  how- 
ever, that  any  additional  material  of  this  rare 
species  will  be  soon  forthcoming,  and  there  is 
little  doubt  that  the  specimen  represents  an  un- 
described  taxon.  The  pelvic  fin  ray  count  of  12, 


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which  I  have  confirmed  on  the  right  fin  by  stain- 
ing, is  distinctly  higher  in  the  eastern  Pacific 
specimen  than  the  9-10  reported  for  E.  mollis 
by  Nybelin  (1957)  and  Marshall  (1973:599).  The 
larger  orbit  (20  percent  HL  cf.  10-15  percent), 
which  goes  into  the  distance  orbit  to  angle  of 
preopercle  about  2.2  times  (compared  with  2.5 
or  more  in  E.  mollis),  the  slightly  more  rays  of 
the  first  dorsal  fin  (11,9  in  E.  occidentalis,  11,10 
or  11,11  in  E.  mollis},  and  the  somewhat  higher 
gill-raker  count  on  the  second  arch  ( 1 1  compared 
with  9-10  for  E.  mollis}  are  other  noteworthy 
differences.  Additionally,  comparison  of  the  ho- 
lotype  of  E.  occidentalis  with  a  specimen  of  E. 
mollis  from  the  Swedish  Deep-Sea  Expedition 
of  1947-48  (reported  on  by  Nybelin  (1957)  and 
borrowed  through  the  courtesy  of  Dr.  Huben- 
dick  of  the  Natural  History  Museum,  Goteborg) 
has  revealed  that  the  scales  on  the  abdomen  of 
the  new  species  are  smaller,  with  about  seven 
scale  rows  separating  the  periproct  from  the  pel- 
vic fin  bases  as  compared  with  three  at  the  most 
separating  the  two  regions  in  E.  mollis.  The  pel- 
vic girdle  of  E.  occidentalis  is  much  shorter  than 
that  of  E.  mollis — its  length  from  anterior  point 
to  base  of  pelvic  fin  goes  about  2.5  into  distance 
orbit  to  angle  of  preopercle,  as  compared  with 
about  2.0  for  the  same  measurement  in  E.  mol- 
lis. The  new  species  is  otherwise  closely  similar 
to  its  congener,  for  which  Nybelin  (1957)  gives 
an  excellent  description  and  a  photograph  of  two 
freshly  caught  specimens. 

DISTRIBUTION. — Known  from  only  the  holo- 
type  taken  off  the  northern  coast  of  Peru  in  4,334 
m. 

MATERIAL  EXAMINED.— Holotype:  USNM  135612  (1,  ca.  80 
mm  HL,  ca.  400  mm  TL);  off  Peru,  8°30'S,  85°36'W,  4,334  m, 
ALBATROSS  sta.  4658,  14  Nov.  1904. 

Mataeocephalus  Berg 

Coelocephalus  GILBERT  AND  CRAMER,  1897:422  (non  Agas- 
siz,  1843)  (type-species  Coelocephalus  acipenserinus  Gil- 
bert and  Cramer,  1897,  by  monotypy). 

Mataeocephalus  BERG,  1898:43  (replacement  name  for  Coe- 
locephalus Gilbert  and  Cramer,  1897,  preoccupied). 

DIAGNOSIS. — Macrourine  grenadiers  with  anus 
located  within  a  moderately  broad  to  broad  na- 
ked area  (periproct)  whose  posterior  edge  is  im- 
mediately adjacent  to  or  close  to  anal  fin  origin; 
anus  usually  closer  to  origin  of  anal  fin  than  to 
insertion  of  pelvic  fins.  Spinous  ray  of  first  dor- 
sal fin  with  a  serrated  leading  edge  or  serrations 
obsolete  (in  M.  adjustus}.  Snout  produced,  dor- 


soventrally  depressed,  with  a  stout,  two-pronged 
scute.  Mouth  small  (less  than  30  percent  HL), 
inferior;  suborbital  ridge  stout,  sharply  angular 
in  cross  section.  Premaxillary  and  mandibular 
teeth  in  broad  bands  which  are  usually  confined 
to  anterior  portion  of  jaws  and  which  usually  fall 
well  short  of  posterior  corners  of  mouth  open- 
ing. Gill  openings  restricted;  gill  membranes 
broadly  connected  to  isthmus.  Outer  gill-rakers 
on  first  arch  rudimentary  or  absent. 

COMPARISONS. — Mataeocephalus  appears  su- 
perficially close  to  Coelorinchus  Giorna  but  is 
immediately  distinguished  from  that  genus  by 
the  branchiostegal  ray  count  of  7  (6  in  Coelorin- 
chus}, the  strongly  serrated  spinous  ray  in  first 
dorsal  fin  (except  in  M.  adjustus  with  serrations 
obsolete;  cf.  smooth  or,  rarely,  with  few  distal 
teeth  in  Coelorinchus},  and  the  two-pronged  ter- 
minal snout  scute  (one-  or  three-pronged  in  Coe- 
lorinchus}. The  genus  Mataeocephalus  is  clos- 
est to  Nezumia  but  differs  in:  (1)  anus  closer  to 
anal  fin  origin  than  to  pelvic  fin  insertion  (anus 
generally  closer  to  pelvic  fin  insertion  in  Nezu- 
mia}; (2)  premaxillary  and  mandibular  teeth  in 
short,  broad  bands  except  in  M.  adjustus  (cf. 
narrow  tapering  bands  that  extend  posteriorly  to 
rictus  except  in  N.  burragei  (Gilbert,  1905));  (3) 
generally  longer  snout,  and  smaller,  more  infe- 
rior mouth;  and  (4)  outer  rakers  of  first  arch  ru- 
dimentary or  absent  (cf.  tubercular  rakers  pres- 
ent in  Nezumia  species).  Members  of  the  genus 
are  unlikely  to  be  confused  with  any  other  ma- 
crourine  grenadier  with  the  combination  of  sev- 
en branchiostegal  rays  and  periproct  adjacent  or 
close  to  anal  fin  origin  because  of  their  small, 
inferior  mouth,  much  produced  snout,  and 
sharp,  angular  suborbital  ridge. 

REMARKS. — Mataeocephalus  includes  a  small 
group  of  about  five  closely  related  species  con- 
fined to  the  tropical  waters  of  the  Pacific  and 
Indian  oceans.  The  genus  is  not  known  from  the 
Atlantic  Ocean. 

Mataeocephalus  adjustus  (Smith  and  Rad- 
cliffe,  1912)  from  the  Philippines  appears  to  be 
the  most  primitive  member  judged  by  the  follow- 
ing characters:  relatively  large  mouth  that  is  not 
strongly  U-shaped;  dentition  in  both  jaws  ex- 
tending posteriad  in  a  tapered  band;  head  and 
snout  not  dorsoventrally  depressed  to  the  extent 
found  in  the  other  members;  ventral  surfaces  of 
snout  and  suborbital  regions  completely  scaled; 
terminal  snout  scute  rather  small;  and  periproct 
area  relatively  small.  Most  of  these  supposedly 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


145 


FIGURE  8.     Mataeocephalus  tenuicauda  (Carman).  Composite  drawing  from  specimens  catalogued  AMNH  8451,  8467,  and 
8468,  collected  by  the  ARCTURUS,  96  km  south  of  Cocos  Island.  Scale  bar  under  otolith  represents  5  mm;  that  under  tail,  25  mm. 


primitive  features  of  M.  adjustus  cloud  the  oth- 
erwise strong  differences  between  the  genera 
Mataeocephalus  and  Nezumia,  but  viewing  the 
group  as  a  whole,  and  on  the  basis  of  the  diag- 
nosis given  above,  recognition  of  each  as  dis- 
tinct genera  seems  justified. 

Mataeocephalus  tenuicauda  (Carman) 
(Figure  8) 

Macrurus  tenuicauda  CARMAN,  1899:216-217.  pi.  49,  fig.  1 
(original  description;  type-locality  Gulf  of  Panama,  838  m, 
ALBATROSS  sta.  3384). 

Mataeocephalus  tenuicauda:  GILBERT  AND  HUBBS,  1916:146 
(name  only). 

DIAGNOSIS. — A  species  of  Mataeocephalus 
with  8  (9  in  three  fins  of  18  specimens)  pelvic  fin 
rays;  22-26  pectoral  fin  rays.  Chin  barbel  about 
4-6  percent  HL.  Upper  jaw  19-28  percent  HL. 
No  small  naked  fossa  (anterior  dermal  window 
of  light  organ)  anterior  to  periproct. 

DESCRIPTION. — General  features  of  fish  seen 
in  Figure  8.  Head  shallow,  depressed;  greatest 
width  of  head  about  equal  to  or  more  than  great- 
est depth  of  head.  Dorsal  and  ventral  surfaces 
of  head  sharply  demarcated  by  a  strong  ridge 


running  from  snout  tip  posteriad  to  preopercle, 
but  not  connected  to  preopercular  ridge.  Mouth 
small,  inferior,  protrusible,  U-shaped.  Periproct 
region  large,  situated  almost  midway  between 
origin  of  anal  fin  and  insertion  of  pelvic  fins,  but 
slightly  closer  to  former.  No  separate  dermal 
window  of  light  organ  apparent  in  specimens 
examined.  Swim  bladder  large,  with  two  long, 
slender,  uncoiled  retia  and  two  small,  flattened 
gas  glands.  Males  with  large  drumming  muscles 
on  each  side  of  anterior  end  of  swim  bladder. 
Pyloric  caeca  simple,  short,  thick;  16  to  21  in 
five  specimens.  Intestine  with  two  major  loops, 
the  first  extending  posterodorsally  from  pylorus, 
the  second  extending  anteroventrally  along  dex- 
tral  wall  of  abdomen,  passing  anteriad  and  si- 
nistrally  around  front  of  stomach  and  caecal 
mass.  Opercular  openings  restricted  dorsally 
and  ventrally;  gill  membranes  broadly  connect- 
ed and  attached  to  isthmus  with  no  free  posterior 
fold.  A  fleshy  ridge  on  shoulder  girdle  along  pos- 
terodorsal  margin  of  gill  cavity.  Outer  gill-rakers 
of  first  arch  rudimentary,  platelike,  3  or  4  in 
number. 

Dorsal  fin  with  thornlike  spinous  first  ray 


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a  l> 

FIGURE  9.  View  of  upper  and  lower  jaws  showing  shape 
of  tooth  bands  in  (a)  Mataeocephalus  tenuicauda  (Garman), 
specimen  48  mm  HL  and  (b)  Nezumia  latirostrata  (Garman), 
specimen  40.5  mm  HL. 


closely  appressed  to  spinous  second  ray,  the  lat- 
ter with  widely  spaced  teeth  along  leading  edge. 
Outer  ray  of  pelvic  fins  prolonged,  extending 
well  beyond  anal  fin  origin. 

Scales  on  body  with  slender,  conical  spinules 
arranged  in  sharp,  ridgelike  rows.  Scales  on 
head  variously  developed;  those  on  ridges  more 
coarsely  developed  with  spinule  rows  arranged 
in  a  stellate  pattern  in  some  and  a  broad  poste- 
riorly radiating  pattern  in  others;  those  on  top 
of  head  generally  with  low,  longitudinal  spinule 
rows  that  diverge  slightly  posteriorly.  Ventral 
surfaces  of  head  naked  except  for  a  small  patch 
of  scales  at  anteroventral  corner  of  preopercle 
and  along  leading  edge  of  snout  where  large, 
coarse,  spinous,  nonimbricate  scales  overlap 
slightly  onto  ventral  surfaces.  Tip  of  snout 
armed  with  two  closely  appressed,  conical 
scutes.  Suborbital  region  covered  dorsally  with 
two  distinct  rows  of  large,  coarse,  nonimbricate, 
strongly  adherent  scales.  Supraorbital  and  su- 
pranarial  ridges  coarsely  scaled.  A  prominent 
lunate  naked  groove  dorsally  along  each  side  of 
anterior  snout  margin. 

Premaxillary  teeth  in  broad,  short,  cardiform 
bands  truncated  at  posterior  ends  (Fig.  9a). 
Mandibular  teeth  in  a  broad,  short  band  with 
tapered  ends;  tooth  band  falling  well  short  of 
lateral  corners  of  mouth. 

Coloration  in  alcohol.  All  specimens  exam- 
ined have  lost  most  of  their  body  scales.  Overall 
color  brown  to  swarthy,  often  with  a  violet  tinge 
on  trunk  and  tail.  Abdominal  region  blackish. 


Naked  ventral  surfaces  of  head  pale  to  dusky. 
Fins  dusky  to  blackish.  Oral,  branchial,  and 
peritoneal  cavities  blackish. 

COUNTS  (from  18  specimens  unless  otherwise 
indicated).— ID.  11,8-10;  IP.  22-26  (x  =  23.76); 
2P.  8-9  (9  rays  in  3  of  36  fins).  Gill-rakers  on 
first  and  second  arch  0-1  +  6-7  (6-8  total;  x  = 
7.18);  on  second  arch  0-1  +  5-7  (6-8  total;  x  = 
7.29).  Scales  below  first  dorsal  7-11  (6  speci- 
mens); below  midbase  of  first  dorsal  6'/i  to  9  (6 
specimens);  below  second  dorsal  SY2  to  11  (9 
specimens);  over  distance  equal  to  predorsal 
length  of  head  37-48  (4  specimens). 

MEASUREMENTS  (from  18  specimens  unless 
otherwise  indicated). — Total  length  131+  to  303 
mm;  head  length  30.0-65.3  mm.  The  following 
in  percent  of  head  length:  postrostral  length  of 
head  60-66  (Jc  =  61.8;  S.D.  =  1.56);  snout 
length  36-42  (x  =  39.9;  S.D.  =  1.46);  preoral 
length  35-43  (x  =  38.6;  S.D.  =  2.31);  internasal 
width  19-22  (x  =  20.5;  S.D.  =  0.80);  orbit  di- 
ameter 26-30  (jf  =  27.2;  S.D.  =  1.48);  interor- 
bital  width  19-22  (x  =  20.3;  S.D.  =  0.93);  post- 
orbital  length  29-36  (Jc  =  32.7;  S.D.  =  1.51); 
orbit  to  angle  of  preopercle  28-33  (x  —  30.0; 
S.D.  =  1.25);  suborbital  width  13-16  (jc  =  14.6; 
S.D.  =  0.96);  upper  jaw  length  19-28  (x  =  22.4; 
S.D.  -  2.07);  barbel  length  3.7-5.7  (x  =  4.7; 
S.D.  =  0.61);  length  outer  gill  slit  8.5-12.7  (x  = 
10.4;  S.D.  -  1.08;  n=  13);  preanal  length  122- 
138  (Jc  =  129.5);  outer  2P.  to  A.  21-29  (Jc  =  25.9; 
S.D.  =  2.16;  /i  =  15);  greatest  body  depth  42- 
56  (*  =  52.2;  S.D.  =  3.47;  n  =  14);  1D.-2D.  in- 
terspace 19-29  (Jc  =  25.1);  height  ID.  52-63  (.v  = 
58.5;  n  =  8);  length  IP.  37^1  (jc  =  38.7;  n  = 
12);  length  2P.  37-49  (x  =  43.2). 

COMPARISONS  AND  RELATIONSHIPS. — Matae- 
ocephalus tenuicauda  closely  resembles  the 
Hawaiian  endemic  M.  acipenserinus  (Gilbert 
and  Cramer,  1897)  but  differs  in  having  a  longer 
upper  jaw  (22-28  percent  HL  vs.  about  19-21.5), 
a  somewhat  longer  chin  barbel  (3.7-5.7  percent 
HL  vs.  2.6-4.0),  and  in  lacking  a  small  naked 
fossa  anterior  to  the  periproct.  M.  nigrescens 
(Smith  and  Radcliffe,  1912)  from  the  Philippines 
also  closely  resembles  both  M.  tenuicauda  and 
M.  acipenserinus,  and  the  three  are  considered 
as  close  allies  by  Gilbert  and  Hubbs  (1920:564). 
My  cursory  examination  of  five  paratypes  of  M. 
nigrescens  (USNM  149310;  149311  [2  speci- 
mens]; 149312;  149313)  revealed  minimal  differ- 
ences between  nigrescens  and  tenuicauda. 
Morphometric  features  of  the  two  are  indistin- 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


147 


guishable.  Scale-row  counts,  however,  show  no- 
table separation;  nigrescens  specimens  had 
about  ll/2  rows  between  the  origin  of  the  second 
dorsal  fin  and  5l/z  below  the  midbase  of  the  first 
dorsal  fin,  whereas  tenuicauda  specimens  had 
Sl/2  to  11  and  6l/2  to  9,  respectively. 

Mataeocephalus  adjustus  (Smith  and  Radcliffe, 
1912)  is  readily  distinguished  from  tenuicauda, 
nigrescens,  and  acipenserinus  by  the  following 
characters:  fewer  pelvic  rays  (7  cf.  8  or  9),  fewer 
pectoral  rays  (19  or  fewer  cf.  21-26),  bands  of 
teeth  in  both  jaws  extending  posteriad  about  to 
end  of  rictus.  Because  of  the  many  features  dis- 
tinguishing M.  adjustus  on  the  one  hand  and  M. 
tenuicauda,  M.  nigrescens,  and  M.  acipenseri- 
nus on  the  other,  a  wide  phylogenetic  diver- 
gence between  the  two  groups  is  suggested.  The 
lack  of  specimens  of  M.  microstomus  (Regan, 
1908)  from  the  Indian  Ocean  and  the  paucity  of 
information  in  the  original  description  preclude 
an  adequate  comparison  of  this  species  with  its 
congeners. 

DISTRIBUTION. — Mainland  Pacific  coasts  of 
Panama  and  Ecuador  between  latitudes  6°36'N 
and  3°15'S;  in  the  Galapagos;  and  south  of  Co- 
cos  Island. 

SIZE. — To  at  least  65  mm  HL  and  303  mm  TL. 

MATERIAL  EXAMINED  (55  specimens,  7  localities). — Pana- 
ma: USNM  148879  (2.  57-59  mm  HL,  202-253  mm  TL), 
6°36'N,  81°45'W,  581  fms  (1,063  m),  ALBATROSS  sta.  4621,  21 
Oct.  1904.  Ecuador:  CAS  38325  (1,  34  HL,  155  TL),  3°15'S, 
80°55'W,  945-960  m,  ANTON  BRUUN  cr.  18B,  sta.  770  (field 
no.  LWK66-120),  10  Sep.  1966.  Cocos  Island  (60  miles  [96 
km]  S  of):  AMNH  8451  (7,  35-64  HL),  AMNH  8467  (6  spec- 
imens), AMNH  8468  (23  specimens),  AMNH  8469  (5  speci- 
mens), AMNH  8470  (7  specimens),  all  from  ARCTURUS  sta. 
74,  May  1925.  Galapagos:  CAS-SU  25239  (1,  55  HL,  230  TL), 
0°29'S,  89°54'30"W,  392  fms  [717  m],  ALBATROSS  sta.  2818,  15 
Apr.  1888.— USNM  135340(1,  43  HL,  185+  TL),  0°36'30"S, 
89°19'00"W,  634  fms  [1,159  m],  ALBATROSS  sta.  2808.  4  Apr. 
1888.— CAS  42075.  (2,  30-31  HL,  158-131+  TL),  1°06'S, 
89°22'W.  700-800  m,  TE  VEGA  cr.  19,  sta.  102,  12  Apr.  1968. 

Paracetonurus  Marshall 

Paracetonurus  MARSHALL,  1973:615  (type-species  Macrurus 
parvipes  Smith  and  Radcliffe,  1912,  by  original  designation). 

DIAGNOSIS. — A  macrourine  grenadier  with 
anus  and  urogenital  openings  within  a  broad, 
black  periproct  immediately  preceding  anal  fin 
origin.  Spinous  ray  of  first  dorsal  fin  with  a  ser- 
rated leading  edge.  Snout  relatively  high,  broad; 
suborbital  region  relatively  deep,  without  a 
strong  spinous  ridge.  Scales  flanking  base  of  sec- 
ond dorsal  fin  not  enlarged.  (Adapted  from  Mar- 
shall 1973.) 


REMARKS. — Marshall  (1973:615)  erected  this 
genus  to  include  Macrurus  flagellicauda  Koe- 
foed,  1927,  M.  parvipes  Smith  and  Radcliffe, 
1912,  and  Lionurus  cetonuropsis  Gilbert  and 
Hubbs,  1916 — three  species  that  are  closely  re- 
lated to  Cetonurus  Giinther,  1887,  but  which  are 
distinguished  by  their  somewhat  less-inflated 
head  and  their  lack  of  enlarged  scales  along  the 
base  of  the  second  dorsal  fin.  The  monotypic 
genus  Kumba  Marshall,  1973,  is  also  closely  re- 
lated to  Paracetonurus  but  differs  primarily  in 
lacking  serrations  on  the  second  spinous  ray  of 
the  first  dorsal  fin. 

I  have  included  Macrurus  fragilis  Garman, 
1899,  in  Paracetonurus  for  reasons  given  in  the 
description  of  that  species.  By  doing  so,  the  def- 
inition of  the  genus  is  expanded  and  its  contrast 
with  the  genera  Cetonurus  and  Kumba  is  less- 
ened. I  recognize,  however,  that  a  detailed  study 
comparing  features  of  P.  fragilis  and  the  three 
other  species  of  the  genus  may  necessitate  a  fur- 
ther rearrangement  of  the  taxon,  but  the  material 
available  does  not  allow  such  a  study  at  this 
time. 

Paracetonurus  fragilis  (Garman) 

(Figure  10) 

Macrurus  fragilis  GARMAN,  1899:203-204,  pi.  46,  fig.  1  (orig- 
inal description;  ALBATROSS  specimens  from  off  Panama 
and  Colombia,  3,058-3,334  m). 

Lionurus  (Lionurus)  fragilis:  GILBERT  AND  HUBBS  1916:146 
(listed). 

Sphagemacrurus  fragilis:  MARSHALL  1973:623  (listed). 

DIAGNOSIS. — A  species  of  Paracetonurus 
with  8-9  pelvic  rays.  Nasal  rostrum  extremely 
fragile;  head  covering  thin,  almost  membranous. 

COUNTS.— ID.  11,6-9;  IP.  19-21;  2P.  8-9. 
Gill-rakers  on  inner  side  of  first  arch  1-2  -I-  8- 
10(10-11  total);  on  inner  side  of  second  arch  1- 
2  +  8-9(9-11  total). 

MEASUREMENTS  (from  three  specimens;  most 
measurements  are  estimates). — Head  length 
about  26-35  mm;  total  length  175+  to  200  mm. 
The  following  in  percent  of  head  length:  snout 
length  about  28^0;  orbit  diameter  about  22-23; 
interorbital  width  24-27;  orbit  to  angle  of  pre- 
opercle  33-44;  suborbital  width  14-18;  length 
upper  jaw  31-39;  length  barbel  19-24;  length 
outer  gill-slit  17-20;  greatest  body  depth  66-69. 

DESCRIPTION. — Head  moderately  wide,  trunk 
short,  tail  very  long  and  straplike.  Head  bones 
thin,  fragile,  particularly  nasal  rostrum,  which 
most  often  is  bent  to  side  giving  appearance  of 


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FIGURE  10.  Paracetonurus  fragilis  (Garman).  (a)  Diagrammatic  reconstruction  from  paratype  MCZ  28585  and  UMML  uncat. 
(b)  SIO  55-265,  snout  reconstructed,  (c)  Stomach  and  intestines  of  UMML  uncat.  (d)  Retia  and  gas  glands  of  UMML  uncat. 
Scale  rule  for  a  and  b  equals  25  mm. 


blunt  snout.  Snout  relatively  high  and  wide.  In- 
terorbital  region  broad,  width  usually  greater 
than  diameter  of  orbit.  Suborbital  region  deep, 
without  a  strongly  angular  ridge.  Mouth  mod- 
erately large,  upper  jaws  extend  to  vertical 
through  hind  margin  of  orbits  or  slightly  beyond. 
Barbel  moderately  long,  slender,  length  equal  to 
or  slightly  greater  than  length  outer  gill-slit.  Gill 
openings  extend  forward  to  below  posterior 
third  of  orbit.  Gill  membranes  little  restricted 
(probably  with  a  moderate  free  fold  posteriorly 
over  isthmus).  Gill-rakers  tubercular,  gill  lamel- 
lae short.  Pyloric  caeca  short,  thick,  10-15  in 
UMML  specimen.  Intestine  short,  only  two 
loops  from  pyloric  caeca  to  anus.  Esophagus 
and  caecal  portion  of  stomach  black;  pyloric 
portion  of  stomach  grayish  brown.  Rectum 
black,  remainder  of  intestine  and  pyloric  caeca 
pallid.  Retia  long,  well  developed,  2;  gas 
glands  2. 

First  dorsal  fin  short,  base  low;  second  spi- 
nous  ray  weakly  serrated  and  produced  beyond 
segmented  rays.  Second  dorsal  fin  rudimentary, 
originating  posterior  to  vertical  through  origin  of 
anal  fin.  Pectoral  and  pelvic  fins  small;  the  latter 
far  forward,  origin  below  preopercle  angle,  with 
outer  ray  slightly  produced  and  extended  past 
origin  of  anal  fin. 

Snout  and  probably  most  of  anterior  and  ven- 


tral parts  of  head  naked  and  covered  with  thin 
black  integument.  (I  could  find  no  scale  pockets 
on  the  head  of  specimens  I  examined,  but  Gar- 
man's  (1899:pl.  46,  fig.  1)  figure  suggests  that 
there  are  some  over  the  gill  covers.)  Garman 
(1899:204)  describes  the  scales  as  "small,  thin, 
deciduous,  with  concentric  striae;  five  scales  in 
a  row  from  the  lateral  line  to  the  base  of  the  first 
dorsal  or  thirty-one  in  a  series  from  this  dorsal 
to  the  anal."  Carman's  figure  of  a  scale  shows 
a  single  point  on  the  posterior  edge,  but  an  oth- 
erwise spinuleless  exposed  field. 

Teeth  short,  slender,  sharp,  in  roughly  two 
rows  in  premaxillae,  in  narrow  band  in  mandi- 
bles; no  enlarged  series. 

Coloration  in  alcohol.  Head  and  abdominal 
region  black,  remainder  of  body  brownish.  Oral, 
branchial,  and  peritoneal  cavities  black.  Gill 
arches  and  rakers  blackish  but  lamellae  pallid. 

DISTRIBUTION. — The  species  is  known  only 
from  the  tropical  eastern  Pacific  where  it  is 
found  in  waters  of  considerable  depths.  Speci- 
mens were  obtained  in  three  trawls  fished  on 
bottom  in  3,058-3,334  m,  and  a  single  specimen 
was  taken  in  an  open  midwater  trawl  fished  to 
a  depth  of  1,335  m. 

COMPARISONS  AND  REMARKS. — As  the  name 
implies,  members  of  this  species  are  extremely 
fragile.  The  few  specimens  available  for  exami- 


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149 


nation  were  in  such  poor  condition  that  an  ad- 
equate description  and  illustration  could  not  be 
prepared.  The  thin,  flexible  rostrum  is  often 
damaged  and  folded  to  the  side  (as  is  the  rostrum 
on  the  specimen  illustrated  by  Garman  1899:pl. 
46,  fig.  1),  but  when  intact,  the  rostrum  is  mod- 
erately long  and  pointed.  The  fragile  head,  and 
particularly  the  rostrum,  is  highly  reminiscent  of 
the  condition  obtaining  in  specimens  of  Hymen- 
ocephalus,  but  members  of  that  genus  and  P. 
fragilis  differ  in  many  other  fundamental  fea- 
tures. The  small  adult  size  of  P.  fragilis  is  in- 
dicated by  the  ripe  condition  of  a  200-mm-TL 
female  paratype  (MCZ  28585),  whose  ovaries 
contained  eggs  as  large  as  1.3  mm  in  diameter. 
This  species  is  quite  unusual  and  may  repre- 
sent a  genus  distinct  from  Paracetonurus,  but 
it  is  placed  in  that  taxon  out  of  convenience, 
because  an  adequate  study  comparing  it  with 
other  related  forms  could  not  be  made.  It  differs 
from  other  members  of  Paracetonurus  in  having 
a  less  inflated  head;  thin,  membranous,  and 
mostly  naked  head  covering  (cf.  moderately 
thick,  completely  scaled  head  covering);  more 
pelvic  fin  rays  (8-9  cf.  6-7);  and  a  larger  mouth 
(upper  jaw  extends  posteriad  to  below  hind  third 
of  orbits,  cf.  below  middle  third  of  orbits).  The 
species  is  also  fairly  close  to  members  of  the 
genus  Sphagemacrurus  Fowler  (in  which  Mar- 
shall (1973)  has  placed  the  species),  but  it  differs 
in  lacking  a  strong  spinous  suborbital  ridge,  a 
high  first  dorsal  fin  base,  and  a  short  blunt 
snout — all  characteristic  of  Sphagemacrurus. 
The  origin  of  the  vent  is  also  more  anteriorly 
placed  in  species  of  Sphagemacrurus  (below 
anterior  third  of  first  dorsal  fin  compared  with 
below  hind  edge  of  first  dorsal  fin  in  P.  fragilis). 

MATERIAL  EXAMINED  (7  specimens.  4  localities). —Pana- 
ma: MCZ  28586  (holotype.  about  230  mm  TL);  6°17'N, 
82°05'W,  3,058  m,  beam  trawl,  ALBATROSS  sta.  3360,  24  Feb. 
1890.  — UMML  uncat.  (1,  about  26  HL,  175+  TL);  6°53'N, 
79°27'W,  3,193  m,  R/V  PILLSBURY  sta.  526,  5  May  1967. 
Colombia:  MCZ  28585  (3  paratypes,  about  29-35  HL,  190+ 
to  200  TL)  and  USNM  57857  (1,  est.  26  HL,  est.  175  TL); 
2°35'N,  83°53'W,  3,334  m,  beam  trawl,  ALBATROSS  sta.  3374, 
3  Mar.  1890.  Eastern  Pacific:  SIO  55-265  (1,  1 17  TL);  00°02'S, 
100°23'W,  0-1,335  m,  3-m  midwater  trawl,  R/V  HORIZON  sta. 
ET(b)-H-65. 

Malacocephalus  Giinther 

Malacocephalus  GUNTHER,  1862:3%  (as  subgenus  of  Macru- 
rus)  (type-species  Macrourus  laevis  Lowe,  1843,  by  mono- 

typy)- 
DIAGNOSIS. — A   macrourine  grenadier  with 


anus  remote  from  anal  fin  and  closer  to  pelvic 
fins;  periproct  large.  Two  large  dermal  windows 
of  light  organ,  the  anterior  one  in  a  bean-shaped 
depression  situated  between  bases  of  pelvic  fins, 
the  posterior  one  in  a  shallow  circular  depres- 
sion close  before  the  anus.  Teeth  large,  widely 
spaced,  in  single  row  in  lower  jaw;  usually  larger 
posteriorly.  Teeth  in  upper  jaw  in  two  rows  or 
in  narrow  band;  outer  series  distinctly  spaced 
and  enlarged.  Pyloric  caeca  numerous  (50-100 
or  more),  multiply  branched.  Lowermost  three 
or  four  branchiostegal  rays  scaled.  Mouth  large, 
upper  jaw  usually  more  than  45  percent  of  head 
length.  No  strongly  developed  scutelike  scales 
on  head  ridges. 

REMARKS. — Malacocephalus  comprises  a 
close-knit  group  of  about  six  species,  one  of 
which  is  undescribed  (see  Iwamoto  1970:410). 
Relationships  of  the  genus  lie  closest  to  Ventri- 
fossa  Gilbert  and  Hubbs,  1920.  a  taxon  repre- 
sented in  the  eastern  Pacific  by  a  single  member 
(of  the  subgenus  Lucigadus),  although  the  genus 
is  common  in  most  other  warm-water  areas. 
Members  of  the  genus  Malacocephalus  are  con- 
fined to  upper-slope  waters  of  tropical  and 
warm-temperate  regions.  Three  of  the  six 
species  (laevis  (Lowe,  1843),  nipponensis  Gil- 
bert and  Hubbs,  1916,  and  hawaiiensis  Gilbert, 
1905)  are  closely  related  and  may  eventually 
prove  to  represent  one  widely  distributed 
species.  Okamura  (1970a:69)  has,  in  fact,  syn- 
onymized  M.  nipponensis  with  M.  laevis. 

Malacocephalus  laevis  (Lowe) 
(Figure  11) 

Macrourus  laevis   LOWE,    1843:92  (original  description;  off 

Madeira). 
Malacocephalus  laevis:  GUNTHER  1862:397-398.  — HUBBS, 

FOLLETT,  AND  DEMPSTER  1979:14  (list;  first  record  from 

eastern  Pacific). 
Macrurus  (Malacocephalus)  laevis:  GUNTHER  1887:148,  pi. 

38,  fig.  b.  See  Marshall  1973:653  for  extensive  synonymy. 
Malacocephalus  sp.:  HUBBS  AND  IWAMOTO  1977:243  (1  spec. 

from  California). 

DIAGNOSIS. — A  species  of  Malacocephalus 
with  spinous  second  ray  of  first  dorsal  fin 
smooth;  teeth  in  upper  jaw  in  two  distinct  rows; 
pectoral  fin  rays  19-20;  upper  jaw  less  than  50 
percent  of  head  length. 

DESCRIPTION  OF  EASTERN  PACIFIC  SPECI- 
MEN.— General  features  of  fish  seen  in  Figure 
11.  Head  compressed  and  deep.  Ridges  not 
sharp  or  coarsely  scaled;  head  contours  smooth- 
ly rounded.  Snout  narrow,  pointed  (in  somewhat 


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FIGURE  11.     Malacocephalus  laevis  (Lowe).  SIO  79-344,  58  mm  HL,  from  off  Santa  Barbara  County,  California,  in  229  m. 


shriveled  eastern  Pacific  specimen;  probably 
rounded  in  life).  Gill  openings  wide,  extending 
anteroventrally  to  below  hind  margin  of  orbit. 
Gill  membranes  loosely  connected  to  isthmus 
with  a  posterior  free  fold.  Interopercle  broadly 
exposed  and  scaled  ventrally.  Periproct  region 
large,  long-ovate,  anus  and  urogenital  opening 
at  broader  posterior  end  of  region;  anterior  der- 
mal window  of  light  organ  in  deep,  lunate  to 
bean-shaped  fossa  between  bases  of  pelvic  fins 
and  separated  from  periproct  by  a  broad  isthmus 
of  scales.  Several  partially  digested  euphausids 
in  stomach.  Pyloric  caeca  fragile,  thin,  branched, 
difficult  to  count,  but  more  than  50  terminal  dig- 
its. 

Scales  uniformly  small  and  finely  spinulated. 
Snout  and  suborbital  region  completely  scaled, 
without  prominent  naked  areas.  Coarse,  scute- 
like  scales  completely  absent;  ridges  of  head  all 


smoothly  rounded.  Lower  jaw  uniformly  scaled. 
Lowermost  branchiostegal  rays  scaled;  gular 
membrane  apparently  without  scales  (but  scales 
here  normally  highly  deciduous  and  may  have 
been  present  in  life). 

Color  gray-brown  overall;  violet  over  abdom- 
inal region,  shoulder  girdle,  and  opercle;  and 
blackish  over  chest,  on  branchiostegal  mem- 
brane, and  on  gular  region  (violet  regions  prob- 
ably not  as  prominent  in  undenuded  and  fresh 
specimens).  Peritoneum  brownish  black.  Oral 
valves  and  lower  lips  peppered  with  melano- 
phores.  remainder  of  oral  cavity  pale.  First  dor- 
sal fin,  pectoral  fin,  pelvic  fin,  and  anterior  por- 
tion of  anal  fin  brownish  black;  second  dorsal 
fin  and  most  of  anal  fin  pale  or  dusky. 

COUNTS.— ID.  11,11;  IP.  20/19;  2P.  9/8.  Gill- 
rakers  on  first  arch  3  +  9;  on  second  arch  3  + 
7.  Scales  below  first  dorsal  approximately  16. 


1WAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


151 


MEASUREMENTS. — Total  length  335  mm;  head 
length  58  mm.  The  following  in  percent  of  head 
length:  postrostral  length  of  head  78;  snout 
length  26;  preoral  length  17;  internasal  width 
17.5;  orbit  diameter  36;  interorbital  width  27; 
postorbital  length  of  head  41;  distance  orbit  to 
angle  of  preopercle  43;  suborbital  width  12;  up- 
per jaw  length  48;  barbel  length  17;  length  first 
gill-slit  26;  preanal  length  123;  outer  pelvic  ray 
to  anal  origin  28;  anus  to  anal  origin  17;  greatest 
body  depth  85;  interspace  between  ID.  and  2D. 
48. 

COMPARISONS  AND  RELATIONSHIPS. — I  have 
tentatively  identified  this  single  eastern  Pacific 
specimen  of  Malacocephalus  as  M.  laevis.  All 
morphometric  and  meristic  characters  examined 
fell  within  the  range  of  variation  enumerated  in 
a  previous  paper  (Iwamoto  1970)  for  specimens 
I  examined  from  the  Atlantic  Ocean.  However, 
the  snout  length  and  interorbital  width  in  the 
Pacific  specimen  were  at  the  lower  limits  of  the 
range  for  these  characters  in  Atlantic  specimens, 
and  the  orbit  diameter  was  at  the  upper  limit. 
Close  comparison  of  the  eastern  Pacific  speci- 
men with  others  from  the  Atlantic  Ocean 
showed  only  one  difference — scales  were  absent 
on  the  gular  membrane  whereas  most  Atlantic 
specimens  had  a  small  patch  of  scales  there.  The 
deciduous  nature  of  the  scales  in  this  area,  how- 
ever, may  account  for  their  absence  in  the  some- 
what denuded  eastern  Pacific  specimen.  Com- 
parison of  SIO  79-344  with  specimens  of  M. 
hawaiiensis  (USNM  51618.  holotype;  CAS-SU 
8522,  3  paratypes)  showed  slightly  lower  values 
in  relative  snout  length  (26  percent  head  length 
cf.  28-29),  preoral  length  (17  cf.  19-21),  inter- 
nasal  width  (17.5  cf.  21-25),  interorbital  width 
(27  cf.  31-35),  length  snout  to  anus  (1 1 1  cf.  117- 
120),  and  1D.-2D.  interspace  (48  cf.  57-75). 
Similar  comparison  of  proportional  measure- 
ments with  specimens  of  M.  nipponensis 
(FAKU  13316,  13318,  13321,  and  13878)  showed 
lower.values  in  the  eastern  Pacific  specimen  in 
snout  length  (26  percent  HL  cf.  28-31),  preoral 
length  (17  cf.  20-21),  interorbital  width  (27  cf. 
30-35),  and  barbel  length  (17  cf.  21-24),  but  a 
greater  value  for  the  orbit  diameter  (36  cf.  29- 
34).  The  gill-raker  count  of  12  was  also  slightly 
higher  than  the  10-11  of  the  M.  nipponensis 
specimens.  Whether  or  not  these  slight  differ- 
ences are  meaningful  in  delimiting  species  is  as 
yet  conjectural.  Adequate  series  of  each  popu- 
lation should  be  compared  before  definitive 


statements  are  made  regarding  the  number  of 
valid  species  in  the  genus.  Until  then  it  seems 
wisest  to  retain  the  established  names. 

REMARKS. — Malacocephalus  laevis  is  listed 
by  Hubbs,  Follett,  and  Dempster  (1979:14)  in 
their  "List  of  the  fishes  of  California"  on  the 
basis  of  the  present  specimen.  Mr.  Eiichi  Fujii 
has  informed  me  (in  litt.,  6  Mar.  1979)  of  having 
collected  specimens  of  the  species  from  sea- 
mounts  off  the  coast  of  Baja  California  while 
aboard  the  KAIYO  MARU  in  January  and  Feb- 
ruary of  1979. 

MATERIAL  EXAMINED.— SIO  79-344  (1,  57.8  mm  HL, 
335  mm  TL).  California,  Santa  Barbara  County,  between  Gav- 
iota  and  Point  Conception,  229  m,  otter  trawl,  10  Dec.  1968. 

Ventrifossa  Gilbert  and  Hubbs 

Ventrifossa  GILBERT  AND  HUBBS,  1920:553  (type-species  Cor- 
yphaenoides  garmani  Jordan  and  Gilbert.  1904,  by  original 
designation). 

DIAGNOSIS. — A  genus  of  macrourine  grena- 
dier with  anus  removed  from  anal  fin  origin  and 
closer  to  pelvic  fin  insertion;  anus  situated  pos- 
teriorly in  an  oval-shaped  area  of  naked  black 
skin  (the  periproct)  that  extends  forward  to  a 
small  fossa  (anterior  dermal  window)  between 
pelvic  fin  bases.  An  often-inconspicuous  lenslike 
posterior  dermal  window  in  front  of  anus.  Sec- 
ond spinous  ray  of  first  dorsal  fin  slightly  pro- 
duced and  finely  toothed  along  leading  edge  (ex- 
cept in  V.  atherodon).  Jaw  teeth  small,  in 
narrow  to  moderately  broad  bands  in  upper  jaw, 
outer  series  slightly  enlarged  in  most  species; 
lower-jaw  teeth  small,  none  enlarged,  in  one  to 
several  irregular  series  laterally.  Branchiostegal 
membranes  naked  (in  subgenus  Ventrifossa)  or 
usually  with  patches  of  scales  (in  subgenus  Lu- 
cigadus)  along  exposed  lowermost  branchioste- 
gal  rays.  No  sharp,  coarsely  scaled,  angular 
ridges  on  head;  suborbital  region  with  flat  to 
gently  rounded  contours — scales  here  not  mod- 
ified to  form  stout  shelves  or  ridges  (as  in  Ne- 
zumia).  Snout  angular  to  rounded  in  profile, 
either  without  a  spiny  terminal  scute  or  with  a 
small  unilateral  one;  supranarial  ridge  without 
modified  scutelike  scales;  lateral  angles  of  snout 
without  spinous  tubercular  scale  at  tip.  Body 
and  head  scales  generally  small,  densely  cov- 
ered with  fine,  slender,  conical  or  shield-shaped 
spinules. 

REMARKS. — Ventrifossa  as  here  diagnosed 
comprises  three  major  groups  which  I  treat  as 
subgenera:  Ventrifossa,  Lucigadus,  and  Soko- 


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dara  (new).  In  describing  Ventrifossa,  Gilbert 
and  Hubbs  (1920:543)  recognized  four  subgen- 
era,  three  of  which  were  monotypic:  Atherodus 
(with  Optonurus  atherodon  Gilbert  and  Cramer, 
1897),  Lucigadella  (with  Macrourus  nigromar- 
ginatus  Smith  and  Radcliffe,  1912),  and  Lugi- 
gadus  (with  Macrourus  lucifer  Smith  and  Rad- 
cliffe, 1912).  Atherodus  was  distinguished  from 
the  other  subgenera  on  the  basis  of  dentition 
(teeth  of  lower  jaw  in  two  series,  arrowhead- 
shaped  canines  on  upper  jaw),  scale  spinules 
(few  and  short),  orbit  size  (3  in  head),  and  dorsal 
spine  (without  denticulations).  My  studies  of  the 
genus  have  shown  that  all  but  the  last  character 
are  nondiagnostic  for  the  subgenus  in  that  the 
character  states  are  shared  with  other  members 
of  the  genus,  or  that  they  are  part  of  a  graduated 
character  spectrum  within  the  genus.  The  ab- 
sence of  denticulations  on  the  dorsal  spine  ap- 
pears to  be  the  sole  character  distinguishing  V. 
atherodon  from  other  members  of  the  genus. 
The  sister-group  relationship  of  V.  atherodon 
and  V.  ctenomelas  (Gilbert  and  Cramer,  1897) 
is  strongly  suggested  in  other  shared  specializa- 
tions and  in  the  common  occurrence  of  the  two 
species  in  the  Hawaiian  Islands,  where  V.  ath- 
erodon is  apparently  endemic  (V.  ctenomelas  is 
also  found  in  the  western  Pacific). 

Characters  that  Gilbert  and  Hubbs  (1920)  used 
to  separate  V.  lucifer  into  a  distinct  subgenus, 
Lucigadus,  include  the  ventral  light  organ  ("a 
conspicuous  pearly  body  in  a  sheath"),  mouth 
angle  ("highly  oblique"),  and  spinules  on  scales 
("in  quincunx  order").  My  reexamination  of  the 
type-specimens  of  V.  lucifer  has  shown  that  the 
"sheath"  in  which  the  "pearly  body"  lies  is  an 
artifact  of  preservation,  the  "sheath"  having 
formed  by  the  ripping  midventrally  of  the  thin 
abdominal  wall  below  the  light  organ.  The  pearly 
color  of  the  organ  is  from  the  reflective  layer 
surrounding  the  organ.  The  light  organ  in  V.  lu- 
cifer is  not  notably  different  from  that  of  other 
closely  related  members  of  the  genus,  including 
V.  nigromarginata.  The  arrangement  of  spinules 
on  the  scales  appears  to  show  no  meaningful 
relationships  in  this  genus  nor  in  the  closely  re- 
lated Nezumia — the  character  is  useful,  how- 
ever, at  the  species  level.  The  notably  oblique 
mouth  of  V.  lucifer  represents  one  extreme  in 
a  graded  series  that  includes — from  mouth  little 
oblique  to  mouth  notably  oblique — Macrourus 
nigromarginatus,  Macrurus  fasciatus  Weber, 


1913,  Macruroplus  ori  Smith,  1968,  Macrourus 
nigromaculatus  McCulloch,  1907,  and  Macrou- 
rus lucifer.  I  consider  these  five  species  as  rep- 
resentatives of  the  subgenus  Lucigadus  Gilbert 
and  Hubbs. 

The  subgenera  of  Ventrifossa  as  I  recognize 
them  can  be  characterized  in  the  following  man- 
ner: 

Subgenus  Ventrifossa  Gilbert  and  Hubbs, 
1920:553  (type-species  Coryphaenoides 
garmani  Jordan  and  Gilbert,  1904). 

1.  Mandibular  teeth  in  one  to  three  irregular 
series  laterally. 

2.  Premaxillary  teeth  in  a  narrow  band  with 
outer  series  slightly  to  prominently  en- 
larged; tooth  band  extends  posteriad  be- 
yond maxillary  process. 

3.  Mouth  slightly  oblique,  moderate  to  large, 
upper  jaw  length  42-53  percent  head  length. 

4.  Tip  of  snout  with  a  blackish  spot,  or  entire 
leading  edge  black. 

5.  Branchiostegal  and  gular  membranes  com- 
pletely naked. 

6.  Gill-rakers  13-20  total  on  inner  series  of  out- 
er (first)  arch. 

7.  Pores  of  cephalic  lateral  line  system  small 
and  inconspicuous. 

8.  Pyloric  caeca  numerous,  40-70  in  distal 
count,  usually  branched  near  base. 

9.  Snout  without  spinous  tubercular  scute  at 
tip. 

10.  Abdominal  vertebrae  11-12. 

1 1 .  Ventral  aspects  of  body  not  appearing  to 
have  shifted  notably  forward. 

12.  Infraorbital  shelf  not  exceedingly  narrow 
anteriorly. 

Included  species:  V.  atherodon  (Gilbert  and 
Cramer,  1897),  V.  ctenomelas  (Gilbert  and  Cra- 
mer, 1897),  V.  diver  gens  Gilbert  and  Hubbs, 
1920,  V.  garmani  (Jordan  and  Gilbert,  1904),  V. 
macropogon  Marshall,  1973,  V.  mucocephalus 
Marshall,  1973,  V.  petersonii  Alcock,  1891,  and 
one  or  two  undescribed  speces  (specimens  in  the 
CAS  collection). 

Subgenus  Lucigadus  Gilbert  and  Hubbs  (type- 
species  Macrourus  lucifer  Smith  and  Rad- 
cliffe, 1912). 

1 .  Mandibular  teeth  in  several  irregular  series 
laterally  or  in  a  narrow  to  moderately  wide 
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153 


2.  Premaxillary  teeth  in  a  narrow  to  moderate- 
ly wide  band  with  outer  series  slightly  en- 
larged; teeth  not  present  posteriad  of  hind 
margin  of  maxillary  process. 

3.  Mouth  slightly  to  greatly  oblique,  moderate 
in  size,  33^5  percent  HL. 

4.  Tip  and  leading  edge  of  snout  without  dis- 
tinct pigmentation. 

5.  Lowermost  of  branchiostegal  rays  scaled. 

6.  Gill-rakers  7-16  total  on  inner  series  of  outer 
(first)  arch. 

7.  Pores  of  cephalic  lateral  line  system  prom- 
inent in  most  species. 

8.  Pyloric  caeca  30-55,  generally  unbranched. 

9.  Snout  tip  without  stout,  spinous,  scutelike 
scale. 

10.  Abdominal  vertebrae  10-11. 

1 1 .  Ventral  aspects  of  body  appearing  to  have 
migrated  far  forward  resulting  in  anal  origin 
below  first  dorsal;  pelvic  origin  below  oper- 
culum;  gill  membranes  united  below  orbits; 
snout  high,  rounded;  base  of  first  dorsal 
high. 

12.  Infraorbital  shelf  not  especially  narrow  an- 
teriorly. 

Included  species:  V.  fasciata  (Weber  and  de 
Beaufort,  1929),  V.  lucifer  (Smith  and  Radcliffe, 
1912),  V.  nigromarginata  (Smith  and  Radcliffe, 
1912),  V.  nigromaculata  (McCulloch,  1907),  and 
V.  ori  (Smith,  1968). 

Subgenus  Sokodara  Iwamoto,  new  subgenus 
(type-species  Coryphaenoides  misakius 
Jordan  and  Gilbert,  1904). 

1 .  Mandibular  teeth  in  a  narrow  band  of  two 
to  three  irregular  series  laterally. 

2.  Premaxillary  teeth  in  a  narrow  band;  outer 
series  scarcely  enlarged. 

3.  Mouth  slightly  oblique,  large,  35-42  percent 
head  length. 

4.  Tip  of  snout  blackish. 

5.  Branchiostegal  and  gular  membranes  com- 
pletely naked. 

6.  Gill-rakers  14-16  total  on  inner  series  of  out- 
er (first)  arch. 

7.  Pores  of  cephalic  lateral  line  system  small 
and  inconspicuous. 

8.  Pyloric  caeca  54-65  in  distal  count,  branched 
near  base. 

9.  Snout  with  a  small,  unilateral,  spinous  scute 
at  tip. 

10.  Abdominal  vertebrae  14. 


11.  Ventral  aspects  of  body  not  appearing  to 
have  shifted  notably  forward. 

12.  Infraorbital  shelf  greatly  narrowed  anterior- 
ly. 

Included  species:  V.  misakia  and  one  or  two 
undescribed  species  (specimens  in  CAS  and 
USNM  collections). 

DISTRIBUTION. — The  genus  Ventrifossa  is 
represented  in  the  eastern  Pacific  by  one  wide- 
ranging,  Southern  Hemisphere  species,  V.  (Lu- 
cigadus) nigromaculata.  That  representatives  of 
the  more  diverse  subgenus  Ventrifossa  are  en- 
tirely absent  from  this  region  forms  a  striking 
parallel  with  a  similar  situation  in  the  eastern 
Atlantic,  where  the  genus  is  entirely  unknown 
(except  off  Cape  Point,  South  Africa) — this  de- 
spite the  presence  in  the  western  Atlantic  of  two 
species  of  subgenus  Ventrifossa  (V.  macropo- 
gon  and  V.  mucocephalus)  and  one  of  subgenus 
Lucigadus  (species  cf.  V.  ori).  Ventrifossa  is 
known  in  most  other  warm-water  regions  of  the 
Pacific  and  Indian  oceans. 

Ventrifossa  nigromaculata  (McCulloch) 

(Figures  \2a,  !&/) 

Macrourm  nigromaculatus  McCutLOCH,  1907:346-348,  pi. 
63,  figs.  1,  la  (original  description;  holotype  and  4  para- 
types;  56  km  E  of  Sydney,  Australia,  in  1,463  m). 

Lionurus  nigromaculatus:  MCCULLOCH  1919:32  (listed),  pi. 
11,  fig.  114a. 

? Macruroplus  nigromaculatus:  SMITH  1949:  135  (brief  de- 
scription; southern  Africa). 

Nezumia  nigromaculata:  MAKUSHOK  1967:207  (name). 

Macruroplus  potronus  Pequeno,  1971:290-291,  fig.  15  (origi- 
nal description;  holotype  only,  off  Chile,  34°58'S,  72°36'W, 
in  200  m). 

DIAGNOSIS. — A  species  of  Ventrifossa,  sub- 
genus  Lucigadus,  with  13-15  pelvic  fin  rays. 
Total  gill-rakers  on  first  arch  12-16;  on  second 
arch  12-14.  Upper  jaw  length  39-45  percent  HL. 
Scales  below  first  dorsal  fin  about  16-20;  below 
second  dorsal  fin  10-12.  First  dorsal  fin  with  a 
prominent  black  blotch. 

DESCRIPTION. — General  features  of  the  fish 
are  best  seen  in  Figure  12a.  Gill  membranes 
unite  across  isthmus  at  a  point  below  middle  of 
orbits  and  slightly  ahead  of  posterior  end  of 
maxillae.  Head  ridges  virtually  nonexistent;  all 
surfaces  smoothly  rounded;  suborbital  region 
almost  flat.  Pores  on  head  well  developed  but 
not  especially  prominent. 

Pyloric  caeca  well  developed,  numerous, 
rather  large;  unbranched  except  at  very  base. 


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Configuration  of  intestinal  tract  a  rather  simple 
'S'  pattern.  Gas  glands  a  flattened  kidney  shape; 
mesial  surfaces  of  the  two  glands  closely  abut- 
ting; retia  attached  mesially  at  about  middle  of 
each  gland.  Retia  slender,  short,  uncoiled. 

Spinules  on  body  scales  very  small,  slender, 
conical,  greatly  reclined,  and  arranged  in  more 
or  less  parallel  horizontal  rows  in  middle  part  of 
vertically  elongate  exposed  field.  Spinule  rows 
number  11-12  on  larger  scales  of  CAS  speci- 
mens. Scales  present  on  lowermost  three  or  four 
branchiostegal  rays  and  also  densely  cover  ex- 
posed ventral  surfaces  of  interopercle. 

Anterior  dermal  window  of  light  organ  large, 
situated  between  pelvic  fin  bases,  and  connected 
to  periproct  by  isthmus  of  naked  black  skin. 
Posterior  dermal  window  relatively  prominent 
in  most  specimens  examined,  discernible  as  a 
discrete  translucent  area  lying  immediately  an- 
teriad  of  anus  and  occupying  most  of  anterior 
portion  of  periproct. 

Fins  well  developed.  First  dorsal  fin  large; 
height  about  equal  to  or  greater  than  length  of 
head.  Anal  fin  high  along  almost  entire  length. 
Pelvic  fins  broad  but  not  especially  long;  outer 
ray  slightly  produced,  extending  slightly  beyond 
anal  fin  origin. 

Coloration  in  alcohol.  Ground  color  tawny  to 
grayish.  Trunk  region  with  violet  tinge  becoming 
bluish  over  abdomen  and  blackish  around  peri- 
proct and  pelvic  fins.  First  dorsal  fin  with  a 
prominent  black  blotch  on  distal  one-half  to  two- 
thirds;  blotch  not  extending  to  margin  of  fin. 
Anal  rays  lightly  punctate  anteriorly  and  black- 
ish along  distal  margins.  Median  fins  blackish 
posteriorly  towards  tail  tip.  Pelvic  fins  blackish 
or  covered  with  bold  punctations.  Gill  covers 
and  gill  membranes  blackish  or  densely  punc- 
tate. Oral  cavity  generally  pale  with  few  scat- 
tered small  areas  blackish  or  densely  punctate. 
Outer  wall  of  gill  cavity  black  along  first  gill-slit 
and  around  outer  margins,  but  pale  otherwise. 
Gill-rakers  and  gill-arches  blackish,  but  gill  fil- 
aments pale.  Lips  dusky  to  somewhat  blackish. 

COUNTS  (from  13  specimens). —  ID.  11,10-11; 


IP.  19-23;  2P.  13-15  (one  specimen  with  12  on 
left  fin,  13  on  right  fin).  Total  gill-rakers  on  first 
arch  12-16;  on  second  arch  12-14.  Scales  below 
first  dorsal  about  16-20;  below  second  dorsal 
10-12;  over  distance  equal  to  predorsal  length 
42-45.  Pyloric  caeca  30,  32,  and  about  57  in 
three  specimens. 

MEASUREMENTS. — Twelve  specimens  exam- 
ined ranged  17-50  mm  HL;  94+  to  339  mm  TL. 
The  following  are  in  percent  of  head  length: 
postrostral  length  73-81  (x=  77.2;  S.D.  =  2.57); 
snout  length  25-30  (x  =  28. 1 ;  S.D.  =  1 .55);  ven- 
tral length  of  snout  18-22  (x  =  19.1;  S.D.  = 
1.38);  orbit  diameter  40-47  (x  =  42.9;  S.D.  = 
2.71);  interorbital  width  20-26  (jf  =  22.1;  S.D.  = 
2.07);  suborbital  width  12-18  (x  =  15.7;  S.D.  = 
2.14);  length  upper  jaw  39-45  (x  =  41.2;  S.D.  = 
1.80);  length  barbel  18-26  (usually  20-26);  length 
outer  gill-slit  23-30  (x  =  26.0;  S.D.  =  1.68); 
preanal  length  140-158;  outer  pelvic  ray  to  anal 
origin  41-54;  anus  to  anal  origin  20-30;  greatest 
body  depth  92-112  (usually  over  100);  depth 
over  anal  origin  75-105;  1D.-2D.  interspace  32- 
58;  height  ID.  101-134;  length  IP.  63-73;  length 
2P.  44-58. 

REMARKS. — Macruroplus  potronus  Pequeno, 
1971,  was  described  from  a  single  specimen  tak- 
en off  Chile  in  200  m.  The  specimen  was  ex- 
amined and  photographed  by  Dr.  William  N. 
Eschmeyer  in  1974.  Examination  of  his  photo- 
graphs and  the  original  description  leaves  no 
doubt  as  to  the  conspecificity  of  that  specimen 
with  those  I  have  examined  from  the  eastern 
Pacific  and  called  Ventrifossa  nigromacnlata. 
Should  these  eastern  Pacific  specimens  prove 
distinct  from  western  Pacific  specimens  now  re- 
ferred to  this  species,  the  name  potronus  re- 
mains available. 

Ventrifossa  nigromacnlata  is  a  distinctive 
species  widely  distributed  in  temperate  waters 
of  the  South  Pacific.  Although  considerable 
variation  was  found  in  several  characters  in  the 
specimens  examined,  the  material  available  did 
not  suggest  a  difference  in  populations  from  op- 
posite sides  of  the  Pacific.  Atlantic  Ocean  spec- 


FIGURE  12.  (a)  Ventrifossa  nigromaculata  (McCulloch).  Specimen  CAS  41668,  35  mm  HL,  from  off  Chile  in  750  m.  (b) 
Nezumia  liolepis  (Gilbert),  CAS-SU  21402.  55  mm  HL,  from  off  Santa  Cruz  Island.  California  in  1,397-1.629  m.  Enlarged  views 
of  scales  from  interorbital  region  and  region  just  below  anterior  end  of  second  dorsal  fin.  (c )  Nezumia  pulchella  (Pequeno), 
CAS  28763.  46  mm  HL,  from  off  Peru  in  272  m.  (d)  Nezumia  stelgidolepis  (Gilbert),  CAS  33109,  56  mm  HL,  off  Pescadero 
Point.  California,  in  439  m. 


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imens  previously  referred  to  this  species  are  ap- 
parently distinct  and  may  represent  an 
undescribed  species. 

DISTRIBUTION. — Australia,  New  Zealand, 
and  Chile,  in  230-1,463  m. 

SIZE. — To  about  50  mm  in  head  length  and 
about  340  mm  total  length. 

MATERIAL  EXAMINED  (13  specimens,  8  localities). — Chile: 
USNM  uncat.  (2,  20-35  mm  HL,  140-246  mm  TL),  NW  of 
Valparaiso,  ca.  400  m;  M/V  Ruiz  I;  11  Feb.  1966.  —CAS 
41669(1,  33  HL,  240  TL),  32°08.5'S,  71°43'W,  960m,  ANTON 
BRUUN  cr.  ISA,  sta.  703  (field  no.  LWK.66-47),  12  Aug.  1966. 
—CAS  41668  (1,  35  HL,  262  TL),  34°06.5'S,  72°18.5'W,  750 
m,  ANTON  BRUUN  cr.  ISA,  sta.  687  (field  no.  LWK66-25),  5 
Aug.  1966. 

New  Zealand:  BMNH  87.12.7.118  (1,  17  HL,  94+  TL), 
CHALLENGER  sta.  166,  275  fms  (503  m).  — LACM  11336-3  (1, 
38  HL,  280  TL),  41°35'S,  175°00'E,  256-490  m,  ELTANIN  sta. 
1848,  19-20  Dec.  1%!.— LACM  10968-12  (5,  25-37  HL,  192- 
283  TL),  44°00'S,  178°06'W,  230-421  m,  ELTANIN  sta.  1398, 
29  Nov.  1964.  — FAKU  42147  (1,  41  HL,  300  TL),  KAIYO 
MARU  sta.  29,  July  1968.  — FSFRL  B3052  (1,  50  HL,  339  TL), 
44°20.5'S,  179°17.5'W,  750  m,  KAIYO  MARU  sta.  33,  15  July 
1968. 

Nezumia  Jordan 

Nezumia  JORDAN,  1904,  in  Jordan  and  Starks:  620  (type- 
species  Nezumia  condylura  Jordan  and  Gilbert,  1904,  by 
original  designation). 

IMacruroplus  BLEEKER,  1874:369  (type-species  Macrourus 
serratus  Lowe,  1843,  by  monotypy)  (nomen  nudum;  see 
Poll  1953:238). 

Lionurus:auct.  (non  Giinther,  1887). 

DIAGNOSIS. — A  macrourine  grenadier  with 
anus  removed  from  anal  fin  origin  (closer  to  pel- 
vic fin  insertion  in  most  species)  and  situated  in 
an  oval-shaped  area  of  naked  black  skin  (the 
periproct).  A  small  fossa  (anterior  dermal  win- 
dow of  ventral  light  organ),  round  to  teardrop 
shaped,  forming  anterior  point  of  periproct  in 
most  species,  detached  from  periproct  in  some 
species.  Teeth  small,  in  narrow  to  broad  bands 
in  both  jaws;  outer  series  of  teeth  in  upper  jaw 
of  most  species  uniformly  spaced  and  slightly 
enlarged.  Teeth  on  premaxillary  bone  do  not 
arise  posteriad  of  maxillary  process.  Mouth 
moderate  to  small,  upper  jaw  length  less  than  40 
percent  of  head  length  in  most  species.  Snout 
pointed  or  bluntly  rounded,  with  stout,  spiny, 
tuberclelike  scales  at  tip  and  lateral  angles, 
prominent  in  almost  all  species.  Barbel  present. 
Suborbital  shelf  covered  with  two  (in  most 
species)  or  more  rows  of  stout,  deeply  embed- 
ded (except  in  N.  liolepis),  nonimbricate,  spi- 
nous  scales;  these  scales  form  a  prominent  edge 
or  crest  that  runs  longitudinally  from  lateral  an- 


gle of  snout  to  preopercle  bone  and  divides  sub- 
orbital  region  into  upper  and  lower  halves.  Body 
scales  covered  with  needlelike  to  shield-shaped 
spinules  (spinules  almost  obsolescent  in  N.  lio- 
lepis).  Snout  and  suborbital  areas  with  some  na- 
ked areas  along  ventral  margins  in  most  species, 
almost  entirely  naked  ventrally  in  these  areas  in 
several  species.  Total  gill-raker  number  on  me- 
sial side  of  outer  (first)  arch  less  than  12  in  all 
but  a  few  species.  Color  various  shades  of  black, 
brown,  blue,  or  violet;  some  species  with  last 
two  colors  have  silvery  reflections  on  body.  Py- 
loric  caeca  unbranched  and  fewer  than  30  in  dis- 
tal count  in  most  species,  as  many  as  60  in  the 
few  species  with  branched  caeca.  Retia  two  (oc- 
casional individuals  may  have  four),  slender, 
uncoiled;  gas  glands  globular,  sometimes  some- 
what flattened. 

REMARKS. — The  genus  Nezumia  is  a  diverse 
group  of  slightly  more  than  40  species  of  which 
ten  are  known  and  confined  to  the  eastern  Pa- 
cific. Most  of  these  ten  species  have  restricted 
distributions,  but  N.  stelgidolepis  is  known  from 
southern  Peru  to  southern  British  Columbia,  and 
N.  convergens  is  common  in  waters  from  north- 
ern Peru  to  Costa  Rica  and  is  here  recorded  from 
the  Islas  Tres  Maria  and  the  Gulf  of  California. 
The  distribution  of  six  species  (latirostrata,  lio- 
lepis,  orbitalis,  parini,  pudens,  pulchella)  each 
spans  fewer  than  30  degrees  of  latitude.  N.  lor- 
icata  is  known  only  from  isolated  captures  in 
the  Galapagos  and  off  central  Chile. 

Although  most  other  genera  are  poorly  rep- 
resented in  the  eastern  Pacific,  the  region  has 
proved  rich  in  Nezumia  species.  The  ten  species 
here  recorded  rank  the  fauna  among  the  largest 
found  in  broadly  comparable  geographical  areas. 
Based  on  my  examination  of  specimens  and 
from  the  literature,  a  breakdown  of  the  number 
of  Nezumia  species  by  area  is  as  follows: 

Eastern  Pacific — 10  spp. 
Atlantic — 14  spp. 

Eastern — 10  spp. 

Western — 9  spp. 

Japan  (8)  +  Philippines  (4) — 10  spp. 
Hawaii — 7  spp. 

Western  Indian  Ocean —  ?4  spp. 
Central  Indian  Ocean —  ?4  spp. 

The  few  species  recorded  from  the  Philippines 
is  surprising,  as  the  area  is  otherwise  exceed- 
ingly rich  in  grenadier  species,  especially  of  the 
genus  Coelorinchus.  Despite  the  extensive  col- 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


157 


lecting  by  the  ALBATROSS  in  the  early  part  of  the 
century,  the  area  remains  poorly  known  in  terms 
of  its  deepwater  fauna,  and  doubtless,  other 
species  of  Nezumia  will  subsequently  be  found 
there.  The  South  China  Sea  and  the  Indo-Aus- 
tralian  Archipelago  likewise  remain  relative  un- 
knowns, and  more  deepwater  collecting  is  badly 
needed  in  these  biologically  rich  areas.  Indian 
Ocean  grenadiers  have  not  been  comprehen- 
sively reviewed  by  recent  workers;  the  generic 
allocations  of  many  species  are  still  question- 
able. 

Members  of  the  genus  are  of  small  to  mod- 
erate size;  only  a  few  attain  lengths  of  more  than 
340  mm  in  total  length  (only  N.  stelgidolepis  in 
the  eastern  Pacific,  but  other  species  in  the  At- 
lantic and  western  Pacific).  The  larger  species 
tend  to  have  a  larger,  more  terminal  mouth  and 
stronger,  longer  teeth — these  features  being 
suggestive  of  a  predatory  habit  on  larger,  more 
active  prey.  The  smaller  species  with  their 
smaller,  more  inferior  mouth,  and  longer,  more 
pointed  snout  most  likely  feed  on  small  prey 
captured  on  or  in  the  bottom  substratum.  Ne- 
zumia parini  is  an  enigma,  however,  in  that  it  is 
strictly  bathypelagic  (the  only  member  of  the 
genus  to  be  so),  yet  it  retains  a  physiognomy 
much  like  that  of  its  bottom-dwelling  congeners 
(see  Hubbs  and  Iwamoto  (1977)  for  additional 
discussion  of  this  peculiar  species). 

The  genus  is  primarily  one  of  upper-  and  mid- 
dle-slope fishes,  with  few  species  ranging  deeper 
than  2,000  m.  Of  the  eastern  Pacific  species 
(Table  2),  N.  convergens  not  only  has  the  sec- 
ond broadest  horizontal  range,  but  also  the 
greatest  vertical  range,  and  it  is  probably  the 
deepest-living  species.  In  contrast,  N.  stelgi- 
dolepis, which  has  the  broadest  horizontal  range 
of  the  eastern  Pacific  species,  has  only  a  mod- 
erate depth  range  and  has  been  captured  at  the 
shallowest  depth. 

The  sole  bathypelagic  member  of  the  genus, 
N.  parini,  has  been  captured  in  midwater  nets 
fished  primarily  in  depths  of  1 ,000  m  or  greater. 
In  the  type-series,  14  of  23  captures  of  the  fish 
were  made  in  nets  fished  to  an  estimated  depth 
of  1,000  m  or  greater;  in  seven,  the  nets  were 
fished  to  depths  estimated  at  between  675  and 
940  m;  and  only  in  two  were  the  nets  fished  shal- 
lower (estimated  depths  of  420  and  455  m) 
(Hubbs  and  Iwamoto  1977). 

Genera  most  closely  related  to  Nezumia  are 
Ventrifossa  Gilbert  and  Hubbs,  1920,  and  Mal- 


TABLE  2.     COMPARISON  OF  CAPTURE  DEPTHS  OF  NINE 
SPECIES  OF  NEZUMIA  FROM  THE  EASTERN  PACIFIC  OCEAN. 

Capture  depths  (m) 


Species 

min. 

max. 

difference 

stelgidolepis 

227 

909 

632 

pulchella 

272 

735 

463 

orbitalis 

523 

800 

277 

ventralis 

549 

717 

168 

pudens 

580 

1,238 

758 

loricata 

600 

1,480 

880 

convergens 

600 

1,865 

1,265 

latirostrata 

605 

1,400 

795 

liolepis 

682 

1,629 

947 

acocephalus  Giinther,  1887;  the  three  constitute 
a  close-knit  unit  which  I  (Iwamoto  1972)  have 
called  the  tribe  Malacocephalini.  The  unifying 
characters  of  this  tribe  are  the  presence  in  the 
members  of  seven  branchiostegal  rays,  a  well- 
developed  periproct  region  that  is  remote  from 
the  anal  fin  origin,  and  the  presence  of  one  or 
two  dermal  windows  of  the  light  organ  anterior 
to  the  anus. 

Nezumia  liolepis  (Gilbert) 

(Figures  \2b,  18c) 

Macrurus  (Lionurus)  liolepis  GILBERT,  1890:117  (original  de- 
scription, off  southern  California,  ALBATROSS  sta.  2980,  in 
1,103  m). 

Lionurus  liolepis:  GOODE  AND  BEAN  1896:409  (occurrence). 

Macrurus  liolepis:  CARMAN  1899:199-200  (description;  rec- 
ords from  ALBATROSS  sta.  3418,  3424,  3436). 

Macrurus  barbiger  GARMAN,  1899: 197,  pi.  45,  figs.  2-2b  (orig- 
inal description;  off  Islas  Tres  Marias,  Mexico,  ALBATROSS 
sta.  3424,  in  676  fm  [1,236  m]). 

Lionurus  (Lionurus)  barbiger:  GILBERT  AND  HUBBS  1916:146 
(listed). 

Lionurus  (Lionurus)  liolepis:  GILBERT  AND  HUBBS  1916:146 
(listed). 

Nezumia  liolepis:  FITCH  AND  LAVENBERG  1968:142  (listed). 

Ventrifossa  barbiger:  MARSHALL  1973:654  (listed). 

DIAGNOSIS. — A  Nezumia  with  10-11  (rarely 
12)  pelvic  fin  rays.  Few  serrations  on  second 
spinous  ray  of  first  dorsal  fin.  Thin,  deciduous 
scales  on  body  with  few  or  no  spinules  on  ex- 
posed field.  Small  ventral  light  organ.  Upper  jaw 
31-40  percent  HL. 

DESCRIPTION. — General  features  of  fish  seen 
in  Figure  12b.  Head  moderately  compressed  and 
deep.  Suborbital  ridge  rounded;  shallow,  obtuse 
angle  formed  by  dorsal  and  ventral  surfaces.  In- 
terorbital  region  shallowly  concave;  width  about 
equal  to  or  less  than  orbit  diameter.  Snout  nar- 
row, bluntly  pointed,  protruding  slightly  beyond 


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rather  large,  almost  lateral  mouth.  Jaws  only 
slightly  restricted  by  lip-folds  at  posterior  angle. 
Opercular  openings  wide,  extending  above  a 
horizontal  through  dorsal  edge  of  orbit;  gill 
membranes  loosely  attached  to  isthmus.  Nape 
high,  strongly  arched  in  large  adults.  First  dorsal 
fin  base  low.  Periproct  region  about  equidistant 
from  pelvic  fin  insertion  and  anal  fin  origin;  ven- 
tral light  organ  poorly  developed,  inconspicuous 
without  dissection.  Gill-rakers  small,  tubercular. 
Gill  filaments  long;  length  of  longest  more  than 
half  diameter  of  orbit,  about  equal  to  or  longer 
than  least  width  of  suborbital  region.  Pyloric 
caeca  slender;  rather  short,  less  than  diameter 
of  orbits;  biramously  branched,  each  main  stem 
branching  one  or  two  times. 

Fin  rays  all  relatively  slender  and  delicate. 
Second  spinous  ray  of  dorsal  fin  slightly  pro- 
longed; leading  edge  smooth  except  for  a  few 
small  denticles  near  distal  end.  Outer  ray  of  pel- 
vic fin  slender;  filamentous  tip  extends  slightly 
beyond  origin  of  anal  fin. 

Overall  coloration  grayish  brown  tinged  with 
violet;  surface  over  abdomen  much  darker,  over 
operculum  blackish,  but  anteriorly  over  rest  of 
head  and  especially  snout  paler.  Lips  dark 
brown  to  blackish;  fins  dusky  to  blackish.  Gill 
membranes  black.  Outer- wall  lining  of  gill  cham- 
ber blackish;  inner-wall  lining  grayish  to  pale. 
Lining  of  oral  cavity  blackish. 

Scales  highly  deciduous;  few  specimens  with 
any  scales  remaining.  A  sizable  patch  remaining 
dorsally  on  head  and  snout  of  CAS-SU  2545,  50 
mm  HL.  Larger  of  these  scales  with  5-7  rows, 
each  with  2-3  long  conical  spinules  (Fig.  12b). 
Scales  over  dorsal  surface  of  snout  either  with 
one  to  several  low  ridges  that  are  sometimes 
armed  with  one  or  a  few  short  spinules,  or  scales 
completely  lack  ridges  and  spinules.  Dorsal  sur- 
face of  snout  lacks  scales  along  leading  edge  and 
along  snout  ridges.  Ventrally,  snout  entirely  na- 
ked back  onto  suborbital  region  to  vertical 
through  end  of  maxillary,  but  a  narrow,  thin 
wedge  of  small,  thin,  cycloid  scales  extends  for- 
ward to  about  level  of  anterior  border  of  orbits. 
Rami  of  lower  jaw  with  small,  fine,  spinuleless 
scales  located  mainly  about  median  axis  of  each 
ramus. 

COUNTS  (from  38  specimens). —  ID.  11,8-11 
(total  10-13;  x  =  11.87;  S.D.  =  0.75);  IP.  20-25 
(x  =  22.72;  n  =  61;  S.D.  -  1.23);  2P.  10-11 
(rarely  12).  Gill-rakers  on  first  arch  1-3  +  7-10 
(total  9-12,  usually  10-11);  on  second  arch 


1-3  +  7-10  (9-12  total).  Scales  below  first  dor- 
sal 8-10;  below  second  dorsal  7-10;  over  dis- 
tance equal  to  predorsal  length  of  head  39-44 
(3  specimens).  Pyloric  caeca  25-37  (6  speci- 
mens). 

MEASUREMENTS  (from  38  specimens). — Total 
length  1 14-290  mm;  head  length  26-63  mm.  The 
following  in  percent  of  head  length  [range  (x; 
n;  S.D.)]:  postrostral  length  of  head  73-81  (77.68; 
34;  1.77);  snout  length  24-28  (25.95;  36;  1.12); 
preoral  length  13-23  (19.12;  34;  2.00);  internasal 
width  17-23  (19.00;  26;  1.50);  orbit  diameter  25- 
31  (27.18;  38;  1.50);  interorbital  width  21- 
27  (23.28;  37;  1.25);  postorbital  length  of  head 
43-56  (50.17;  35;  3.42);  distance  orbit  to  angle 
of  preopercle  36-44  (39.39;  37;  1.88);  suborbital 
width  11-15  (12.92;  37;  0.96);  upper  jaw  length 
(28)  31-40  (34.12;  38;  2.31);  barbel  length  10-20 
(14.68;  37;  2.59);  length  first  gill-slit  14-23 
(17. 38;  37;  1.65);  preanal  length  119-142(131.52; 
27;  5.81);  outer  pelvic  ray  to  anal  origin  32-57 
(42.26;  26;  6.24);  anus  to  anal  origin  13-27 
(19.76;  24;  4.66);  greatest  body  depth  60-82 
(70.89;  29;  4.86);  1D.-2D.  interspace  28-48 
(34.63;  36;  6.15);  ID.  height  58-73  (62.267;  21; 
4.27);  IP.  length  43-53  (49.15;  24;  3.30);  2P. 
length' 42-55  (48.03;  25;  6.13). 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
wrnia  liolepis  belongs  with  that  group  of  Ne- 
zumia  species  characterized  by  (1)  a  moderate- 
sized  mouth,  (2)  a  relatively  deep,  compressed 
head,  (3)  a  rather  blunt,  high,  and  usually  weakly 
armed  snout,  and  (4)  relatively  wide  gill  open- 
ings. This  group  includes  such  geographically 
separated  species  as  N.  stelgidolepis  (Gilbert), 
N.  atlantica  (Parr,  1946),  N.  africana  (Iwamo- 
to,  1970),  N.  bubonis  Iwamoto,  1974,  N.  dara 
(Gilbert  and  Hubbs,  1916),  N.  burragei  (Gilbert, 
1905),  N.  hebetata  (Gilbert,  1905),  N.  holocen- 
trus  (Gilbert  and  Cramer,  1897),  N.  kamoharai 
Okamura,  1970,  and  N.  macronema  (Smith  and 
Radcliffe,  1912).  N.  liolepis  is  readily  distin- 
guished from  these  species  in  having  the  com- 
bination of  (1)  few  serrations  on  the  dorsal  spine, 
(2)  reduced  spinulation  on  the  scales  of  the  head 
and  body.  (3)  extensive  naked  areas  on  the  dor- 
sal and  ventral  surfaces  of  the  snout  and  sub- 
orbital  region,  (4)  a  distinctive  shape  of  the  oper- 
culum, and  (5)  a  relatively  posteriad  position  of 
the  anus. 

REMARKS.— Garman  (1899:197)  described 
Macrurus  barbiger  from  a  10-inch  (25-cm)  spec- 
imen taken  off  the  state  of  Guerrero,  Mexico. 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


159 


He  contrasted  the  species  with  /V.  liolepis,  re- 
porting that  barbiger  has  "the  head  more  round, 
the  cheeks  more  convex,  the  snout  narrower, 
and  the  barbel  longer,  besides  which  differences 
there  are  others  in  the  fins  and  the  colors."  I 
compared  the  holotype  of  M.  barbiger  (MCZ) 
28597)  with  specimens  Garman  identified  as 
"Macrurus  liolepis"  and  found  that  these  dif- 
ferences do  not  hold  up.  There  is  little  doubt 
that  the  M.  barbiger  holotype  and  the  "M.  lio- 
lepis" specimens  are  conspecific. 

DISTRIBUTION. — The  species  is  known  from 
off  Monterey  Bay.  California  (36°49'20°N),  to 
south  of  Guerrero,  Mexico  (17°24'N),  including 
the  Gulf  of  California  south  of  Guaymas  (Gar- 
man 1899:200.  ALBATROSS  sta.  3436).  Capture 
depths  have  ranged  from  768-823  m  (CAS 
26638)  to  1.655  m  (ALBATROSS  sta.  3436). 

MATERIAL  EXAMINED  (40  specimens  from  9  localities). — 
California  (north  to  south):  CAS-SU  5351  (4,  28-52  mm  HL), 
off  Monterey  Bay.  36°49'20"N,  122°12'30"W,  834  m,  ALBA- 
TROSS sta.  3126,  13  Mar.  1889.  —CAS-SU  21402  (5.  53-63). 
off  Santa  Cruz  Islands,  1.397-1,629  m,  ALBATROSS  sta.  4428. 
14  Apr.  1904.  — USNM  44271.  syntypes  (2,  45-50),  E  of  Santa 
Cruz  Islands,  33°49'45"N,  119°24'30"W,  1,103  m,  ALBATROSS 
sta.  2980,  12  Feb.  1889.  —CAS  26638  (4,  41-56),  off  San  Ma- 
teo  Point,  33°15'30"N,  117°38'W,  768-823  m,  N.  B.  SCOFIELD 
sta.  53B59,  23  June  1953.  —CAS-SU  2545  (6,  27-50),  off  San 
Diego.  32°49'N,  117°27'30"W,  656  m,  ALBATROSS  sta.  2936.  4 
Feb.  1889.  —USNM  77495  (3,  33-33),  off  Point  Loma.  San 
Diego.  940-989  m,  ALBATROSS  sta.  4333,  9  Mar.  1904. 

Mexico:  MCZ  28597  (holotype  of  Macrurus  barbiger  Gar- 
man. 1899.  54  mm  HL),  off  Islas  Tres  Marias,  21°15'N, 
106°23'W,  1,236  m,  ALBATROSS  sta.  3424,  18  Apr.  1891.  - 
CAS  40230  (10,  38-47),  off  Jalisco,  19°43.5'N,  105°35.5'W, 
700-900  m.  TE  VEGA  cr.  19,  sta.  17.  10  July  1968.  —CAS 
40231  (5,  51-58).  off  Guerrero,  17°24'N,  10P31'W,  940-1,000 
m.  TE  VEGA  cr.  19.  sta.  19.  12  July,  1968. 

Nezumia  pulchella  (Pequeno,  1971) 

(Figures  12c,  ISb) 

Macruroplus  pulchellus  PEQUENO,  1971:293-294,  fig.  17  (orig- 
inal description;  off  Chile;  type-locality  25°26'S.  70°37'W, 
374^*24  m). 

DIAGNOSIS. — A  species  of  Nezumia  with  11- 
12  pelvic  fin  rays.  Gill-rakers  on  first  arch  9-11 
total;  on  second  arch  8-10.  Spinules  on  body 
scales  conical,  in  distinct,  slightly  convergent 
rows.  Ventral  surfaces  of  snout,  suborbital  re- 
gion, mandibular  rami,  and  gular  and  branchio- 
stegal  membranes  usually  naked,  but  occasion- 
ally with  few  scattered  scales.  Barbel  9-16 
percent  of  HL;  upper  jaw  30-34  percent  HL. 

DESCRIPTION. — General  features  offish  seen 
in  Figure  12c.  Head  moderately  compressed, 
trunk  deep  in  large  adults,  less  so  in  smaller  in- 


dividuals. Suborbital  ridge  prominent  in  study 
material,  accentuated  by  shrunken  lower  portion 
of  region;  suborbital  ridge  of  fresh  specimens 
probably  not  as  pronounced.  Other  head  ridges 
rather  smoothly  rounded  without  reinforcing 
spiny  scutelike  scales.  Snout  narrow  and  short, 
relatively  blunt;  armed  with  small  spiny  scutes 
at  tip  and  at  lateral  angles.  Mouth  moderately 
large,  lateral,  unrestricted  by  lip  folds  at  angle. 
Posterodorsal  corner  of  opercle  slightly  pro- 
duced. Interopercle  broadly  exposed  posteriorly 
and  ventrally.  Gill  openings  wide,  membranes 
broadly  connected  over  isthmus  with  a  moder- 
ately broad  free  fold.  Anterior  dermal  window 
of  light  organ  small,  in  shallow  fossa  between 
bases  of  pelvic  fins  and  slightly  removed  from 
periproct.  Gill  filaments  long;  length  of  longest 
about  equal  to  diameter  of  pupil,  greater  than 
width  of  suborbital  region.  Pyloric  caeca  well 
developed,  but  fairly  short  (about  0.6  into  orbit 
diameter),  branched  only  at  bases;  23-31  in  nine 
specimens. 

Scales  of  body  with  distinct,  slightly  conver- 
gent rows  of  small  conical  spinules  on  exposed 
fields.  Scales  generally  lacking  on  gular  mem- 
brane, mandibles,  lower  part  of  snout  and  sub- 
orbital  region,  and  branchiostegal  membrane, 
but  one  specimen  (CAS  38323)  with  two  small 
scales  on  right  gill  membrane  near  base  of  fifth 
branchiostegal  ray.  Small  areas  dorsally  behind 
leading  edge  of  snout  naked.  Stout,  spiny  scales 
at  tip  and  lateral  angles  of  snout  and  in  two  rows 
along  dorsal  surface  of  suborbital  region,  but 
scales  otherwise  not  modified.  Sensory  pores 
along  ventral  margins  of  suborbital  region  and 
preopercle,  and  along  inner  margins  of  mandi- 
bles large  and  prominent. 

Teeth  in  both  jaws  in  rather  narrow  band; 
teeth  small  except  for  outer  premaxillary  series, 
which  is  composed  of  enlarged,  spaced,  sharp, 
conical  teeth. 

Coloration  in  alcohol  medium  brown  overall 
with  bluish  tinge  over  abdomen  and  blackish 
ventrally  on  trunk.  Gill  membranes  black;  fins 
blackish.  Lining  of  buccal  cavity  pale  except  for 
grayish  oral  valves.  Peritoneal  cavity  pale  but 
peppered  with  small  melanophores. 

COUNTS  (from  16  specimens). —  ID.  11,10-11 
(9  in  one  specimen);  IP.  19-28  (x  =  24.52;  n  = 
31;  S.D.  -  2.06);  2P.  11-12.  Mesial  gill-rakers 
on  first  arch  1-3  +  7-9  (9-11  total);  on  second 
arch  1-2  +  7-9  (8-10  total).  Scales  below  first 
dorsal  61/i-81/i;  below  second  dorsal  SVi-lVi; 


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below  midbase  of  first  dorsal  4Vi-6;  over  dis- 
tance equal  to  predorsal  length  35^44  (usually 
37-40). 

MEASUREMENTS  (from  15  specimens,  22-mm- 
HL  specimen  from  SIO  65-675  excluded). — To- 
tal length  137+  to  286+  mm;  head  length  29-61 
mm.  The  following  in  percent  of  head  length 
[range  (x;  n\  S.D.)]:  postrostral  length  of  head 
79-82  (80.47;  15;  1.06);  snout  length  21-26 
(23.59;  15;  1.38);  preoral  length  16-19(17.61;  15; 
0.62);  internasal  width  15-21  (18.47;  15;  1.42); 
orbit  diameter  30-34  (32.29;  15;  1.47);  interor- 
bital  width  20-25  (22.79;  15;  1.31);  postorbital 
length  40^6  (43.34;  15;  1.88);  orbit  to  angle  of 
preopercle  34-38  (36.05;  15;  1.30);  suborbital 
width  11-13  (11.85;  15;  0.66);  upper  jaw  length 
30-34  (32.15;  15;  1.29);  length  barbel  9-16 
(11.73;  15;  3.38);  length  first  gill-slit  17-23 
(19.95;  15;  1.54);  preanal  length  117-150(131.93; 
15;  7.51);  greatest  body  depth  68-88  (79.80;  15; 
5.43);  1D.-2D.  interspace  31-56  (41.83;  15; 
7.40);  height  first  dorsal  fin  65-76  (69.30;  10; 
4.00);  length  pectoral  fin  51-70(61.53;  15;  5.17); 
length  pelvic  fin  39-53  (45.87;  15;  3.48). 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
zumia pulchella  is  closely  related  to  N.  stelgi- 
dolepis  but  differs  primarily  in  having  11-12 
pelvic  fin  rays  rather  than  the  9-10  of  N. 
stelgidolepis.  Scale  spinules  also  show  notable 
differences  between  the  two  species.  In  N.  pul- 
chella the  spinules  are  all  conical  and  in  distinct 
rows  that  converge  slightly  towards  the  midline, 
whereas  in  stelgidolepis  the  spinules  are  conical 
to  lanceolate  and  arranged  in  a  more  quincunx 
pattern.  Retia  and  gas  glands  of  pulchella  are 
also  much  larger  and  stouter,  the  barbel  is  short- 
er, and  the  branchiostegal  rays  are  naked  except 
for  occasional  scattered  scales  (as  opposed  to 
short  rows  of  deciduous  scales  along  the  bases 
of  the  lowermost  branchiostegal  rays  in  stelgi- 
dolepis). 

Nezumia  pulchella  is  readily  distinguishable 
from  N.  pudens  in  having  extensive  naked  areas 
on  the  ventral  surface  of  the  head  and  on  the  man- 
dibles (as  compared  with  surfaces  mostly  scaled 
in  pudens),  fewer  scale  rows  below  the  first  and 
second  dorsal  fins  (6l/2  to  Sl/2  vs.  lOVi  to  12  be- 
low the  first  dorsal;  5!/2  to  71/2  vs.  8»/2  to  W/2 
below  the  second  dorsal,  fewer  pyloric  caeca 
(23-31  vs.  37-44),  and  a  shorter  barbel  (9-16  vs. 
20-25). 

SIZE. — To  at  least  286  mm  in  total  length  (in 


a  specimen,  IMARPE  uncat.,  61  mm  in  head 
length).  A  specimen  in  the  type-series  measured 
67  mm  HL  and  272  mm  TL  (Pequeno  1971:294, 
table). 

DISTRIBUTION. — Northern  Peru  (7°49'S)  to 
central  Chile  (about  33°S)  in  250-960  m. 

MATERIAL  EXAMINED  (16  specimens  from  8  localities). — 
Peru:  CAS  38323  (3,  54-57  mm  HL,  207+  to  270  mm  TL), 
7°49'S,  80°38'W,  605-735  m,  ANTON  BRUUN  cr.  18B,  sta.  754 
(field  no.  LWK66-93),  5  Sep.  1966.  —CAS  28761  (1,  43  HL, 
187  TL)  and  IMARPE  uncat.  (1,  61  HL,  286+  TL),  13°49.4'S, 
76°46.9'W,  570  m,  24  Jan.  1972.  —CAS  28763  (1,  46  HL,  226 
TL),  13°53.3'S,  76°42.0'W,  272  m,  25  Jan.  1972.  —CAS  28764 
(1,  35  HL,  164  TL),  16°31.0'S,  73°12.2'W,  510  m,  1  Feb.  1972. 
—IMARPE  uncat.  (1,  34  HL,  161  TL),  18°07'S,  71°02.5'W,  28 
Jan.  1972.  Chile:  CAS  38320  (6,  26-39  HL,  135-185  TL), 
23°41'S,  70°34'W,  250-400  m,  ANTON  BRUUN  cr.  ISA,  sta. 
717  (field  no.  LWK66-63),  17  Aug.  1966.  —CAS  38318  (1,  38 
HL,  158  TL),  32°08.5'S,  71°43'W,  960  m,  ANTON  BRUUN  cr. 
18A,  sta.  703  (field  no.  LWK66-47),  12  Aug.  1966.  —SIO  65- 
675  (1,  22  HL,  104  TL),  about  29  km  off  Valparaiso  Harbor 
(ca.  33°N),  22-23  Dec.  1965. 

Nezumia  stelgidolepis  (Gilbert) 
(Figures  \1d,  13,  18i) 

Macrurus  stelgidolepis  GILBERT,  1890:116  (original  descrip- 
tion; off  Pt.  Conception,  California,  ALBATROSS  sta.  2960, 
in  488  m). 

Macrurus  gracillicauda  GARMAN,  1899:206-207,  pi.  H,  fig.  1 
(original  description;  Gulf  of  Panama,  ALBATROSS  sta.  3384 
and  3385,  in  837  and  523  m). 

Lionurus  (Nezumia)  stelgidolepis:  GILBERT  AND  HUBBS 
1916: 145  (listed). 

Lionurus  stelgidolepis:  BARNHART  1936:24  (brief  description). 

Nezumia  stelgidolepis:  ROEDEL  1951:509,  fig.  183  (16  records 
off  California). 

Nezumia  gracillicauda:  MARSHALL  AND  IWAMOTO  in  MAR- 
SHALL 1973:626  (listed). 

DIAGNOSIS. — A  species  of  Nezumia  with  9- 
10  pelvic  fin  rays.  Gill-rakers  on  first  arch  8-12 
total;  on  second  arch  8-11  total.  Spinules  on 
body  scales  conical  to  narrowly  lanceolate.  Ven- 
tral surfaces  of  snout,  most  of  suborbital  region, 
and  anterior  half  or  more  of  mandibular  rami 
naked.  Lowermost  branchiostegal  rays  scaled. 
Barbel  15-26  percent  HL;  upper  jaw  31-37  per- 
cent. 

DESCRIPTION. — General  features  of  fish  seen 
in  Figures  12d  and  13.  Head  moderately  com- 
pressed and  deep;  ridges  of  head  not  strongly 
developed,  usually  somewhat  rounded;  subor- 
bital region  relatively  flat.  Snout  narrow,  bluntly 
pointed  (more  pointed  in  young),  protruding 
slightly  beyond  rather  large  mouth,  which  is 
only  slightly  restricted  at  posterior  angle  by  lip 
folds.  Posterodorsal  corner  of  opercle  somewhat 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


161 


a 


FIGURE  13.  Nezumia  stelgidolepis  (Gilbert),  (a)  CAS  44197.  14.7  mm  HL,  73  mm  TL  (fins  and  head  region  somewhat 
reconstructed  after  13.5-mm-HL  specimen  from  same  station).  (ft)  USNM  120272,  28.7  mm  HL,  280  mm  TL,  partially  recon- 
structed, (c)  CAS  40022,  53  mm  HL,  255  mm  TL. 


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flaplike,  posterior  margin  of  gill  cover  steeply 
oblique  along  opercle,  gently  rounded  along  sub- 
opercle,  connecting  to  broadly  exposed  (and 
scaled)  interopercle.  Shape  of  gill  cover  similar 
to  that  in  N.  liolepis.  Gill  openings  wide;  gill 
membranes  rather  narrowly  connected  over 
isthmus,  with  free  posterior  fold.  Nape  region 
slightly  arched.  Anus  within  lenticular  area  of 
black  naked  skin,  situated  slightly  behind  bases 
of  pelvic  fins,  but  closer  to  these  than  to  anal 
fin.  Anterior  dermal  window  of  light  organ  small, 
in  shallow  fossa  between  pelvic  fin  bases.  Gill 
filaments  well  developed;  length  longest  filament 
equal  to  or  longer  than  least  suborbital  width 
and  about  equal  to  pupil  diameter.  Pyloric  caeca 
numerous,  branched;  counts  highly  variable,  in 
eight  specimens  24,  34,  36,  38,  38,  40,  43,  and 
58.  Periproct  region  relatively  large,  close  be- 
hind pelvic  fin  bases;  anterior  window  of  light 
organ  small,  circular,  lying  between  inner  edges 
of  pelvic  fin  bases. 

Scales  densely  covered  with  long,  narrowly 
lanceolate  to  conical  spinules  arranged  in  short, 
slightly  convergent  rows.  Ventral  surfaces  of 
snout  and  most  of  suborbital  region,  and  anterior 
half  to  two-thirds  of  lower  jaw  naked.  Sensory 
pores  rather  prominent  over  these  naked  areas. 
Short  rows  of  small  deciduous  scales  along  bas- 
es of  one  or  more  branchiostegal  ray. 

Coloration  of  fresh  specimen  (CAS  40022) 
swarthy  overall  with  blackish  predominant  over 
ventral  aspects  of  abdomen,  chest,  head,  and  gill 
covers.  Bluish  tinge  over  blackish  abdominal 
walls.  Fins  dusky  to  blackish;  first  dorsal  slightly 
paler  basally.  Lining  of  buccal  cavity  pale,  but 
oral  valves  gray.  Pharyngeal  region  dark  gray; 
gill-rakers  gray  but  gill  filaments  pale.  In  juve- 
niles (CAS  41 196  and  41 197),  oral  cavity  densely 
peppered  with  small  melanophores. 

Dentition  in  broad  bands  in  both  jaws,  nar- 
rowly tapered  posterolaterally.  Outer  series  of 
upper  jaw  teeth  slightly  enlarged  and  evenly 
spaced. 

Four  juvenile  specimens  presumably  of  this 
species  were  captured  in  midwaters.  The  largest 
of  these  (LACM  30611-16)  measured  19  mm  in 
HL,  106  mm  in  TL,  and  was  taken  in  an  Isaacs- 
Kidd  midwater  trawl  off  the  northwestern  coast 
of  Baja  California  between  the  surface  and  about 
600  m,  over  a  bottom  depth  of  1,600  to  1,100 
fms  (2,962-2,012  m).  This  specimen  was  dark 
colored,  superficially  resembling  N.  parini,  but 
the  coarse  scale  spinules,  the  large  pores  on  the 


head  and  mandibles,  the  distinct  lateral  line,  the 
number  of  pelvic  fin  rays,  and  the  large  gill  fil- 
aments distinguish  the  species  as  N.  stelgido- 
lepis.  The  three  other  specimens  (CAS  41196 
and  41 197)  captured  in  midwaters  were  too  small 
to  positively  identify  without  more  comparative 
material,  but  the  general  appearance  (see  Fig. 
13c)  and  counts  (of  dorsal  and  pelvic  fin  rays, 
and  gill-rakers)  also  suggest  N.  stelgidolepis  as 
the  correct  identification.  Proportional  measure- 
ments of  these  juveniles  were  generally  diver- 
gent from  those  of  the  adults,  indicating  sub- 
stantial allometric  growth  in  such  features  as 
preoral  length  of  snout,  orbit  diameter,  inter- 
obital  width,  postorbital  length  of  head,  distance 
orbit  to  angle  of  preopercle,  upper  jaw  length, 
and  barbel  length. 

The  smallest  examined  specimen  (USNM 
57861)  captured  in  a  bottom  trawl  measured  39 
mm  in  head  length  and  176  mm  in  total  length. 

COUNTS  (from  33  specimens). —  ID.  11,9-10 
(8  in  one  specimen,  11  in  two  specimens);  IP. 
20-26  (x  =  21.00;  n  =  59;  S.D.  =  3.30);  2P.  9- 

10  (8  in  one  specimen,  11  in  two  others).  Gill- 
rakers  on  first  arch  10-11  total  (8  in  one,  9  in 
one,  12  in  three  specimens);  on  second  arch  9- 

1 1  total.  Scales  below  first  dorsal  8-9  (rarely  10); 
below  second  dorsal  7-8  (9  in  one);  below  mid- 
base  of  first  dorsal  6-7;  over  distance  equal  to 
predorsal  length  of  head  34—47,  usually  between 
36  and  42. 

MEASUREMENTS  (from  38  specimens,  juve- 
niles excluded). — Total  length  176-405+  mm; 
head  length  39-96  mm.  The  following  in  percent 
of  head  length  [range  (x;  n;  S.D.)]:  postrostral 
length  of  head  73-82  (78.60;  35;  1.80);  snout 
length  22-28  (24.51;  36;  1.34);  preoral  length  12- 
18  (14.86;  36;  1.28);  internasal  width  14-19 
(16.46;  30;  1.19);  orbit  diameter  26-32  (28.06;  38; 
1.68);  interorbital  width  20-26  (23.46;  38;  1.55); 
postorbital  length  43-52  (46.56;  33;  2.17);  orbit 
to  angle  of  preopercle  35-45  (39.81;  37;  2.01); 
suborbital  width  11-16  (12.84;  37;  1.01);  upper 
jaw  length  31-37  (34.93;  38;  1.49);  length  barbel 
15-26  (20.98;  36;  2.72);  length  first  gill-slit  12-18 
(16.37;  36;  1.54);  preanal  length  119-157(135.95; 
37;  9.51);  greatest  body  depth  70-90  (77.68;  33; 
5.77);  1D.-2D.  interspace  26-58  (45.92;  38; 
11.19);  height  first  dorsal  fin  48-68  (60.73;  29; 
4.26);  length  pectoral  fin  44-56  (49.69;  35;  3.23); 
length  pelvic  fin  35-49  (43.48;  34;  3.61). 

COMPARISONS  AND  RELATIONSHIPS. — In  the 
eastern  Pacific,  Nezumia  stelgidolepis  is  most 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


163 


closely  related  to  N.  liolepis  and  N.  pulchella, 
but  is  readily  distinguished  from  these  two  by 
differences  in  scale  spinulation,  squamation  pat- 
tern, pelvic  ray  count,  and  other  features  noted 
in  the  key.  Nezumia  stelgidolepis  is  represen- 
tative of  that  group  of  Nezumia  spp.  character- 
ized by  relatively  blunt  snout,  large  mouth, 
broadly  unrestricted  gill  openings,  deep  body, 
and  large  size.  Some  of  the  species  in  this  group 
(e.g.,  Nezumia  atlantica  (Parr,  1946),  N.  afri- 
cana  (Iwamoto,  1970))  have  been  treated  in  oth- 
er works  (Parr  1946;  Iwamoto  1970;  Marshall 
1973)  as  members  of  the  genus  Ventrifossa. 

SIZE. — Nezumia  stelgidolepis  is  the  largest 
eastern  Pacific  member  of  the  genus,  attaining 
a  total  length  of  at  least  445  mm  (see  Roedel 
1951:509).  Nezumia  atlantica,  a  close  relative 
from  the  western  Atlantic,  attains  a  comparable 
size,  probably  exceeded  in  the  genus  only  by  the 
peculiar  species  N.  bubonis  Iwamoto,  1974. 
from  the  Hawaiian  Islands  and  the  western  At- 
lantic. 

DISTRIBUTION. — British  Columbia  (off  Van- 
couver Island)  to  southern  Peru  (18°10.0'S),  in 
277-909  m  (Makushok  (1967:table  18)  gives  the 
depth  distribution  as  "(61-91)379-909"). 

MATERIAL  EXAMINED  (excluding  material  previously  listed 
in  Iwamoto  and  Stein  1974:49-50).— California:  CAS  233% 
(1,  63  mm  HL,  310  mm  TL),  off  Ft.  Bragg  in  475-494  m.  — 
CAS  33110  (1,  43  HL,  243  TL).  off  Bodega  Bay,  494-585  m, 
23  Feb.  1965.  —CAS  31509  (1.  73  HL.  365+  TL).  off  Point 
San  Simeon,  530  m,  N.  B.  SCOFIELD  sta.  54BB71,  25  Oct. 
1954.  —CAS  40022  (1,  53  HL,  255  TL),  off  Santa  Barbara 
County,  34°18.4'N,  120°14.4'W,  439^43  m.  COMMANDO,  10 
Aug.  1977.  —  USNM  87579  (1 ,  68  HL).  ALBATROSS.  — USNM 
127072  (1.  39  HL,  280+  TL),  off  southern  California,  ALBA- 
TROSS. 

Mexico:  LACM  30611-16  (1,  19  HL,  106  TL),  Guadalupe 
I.,  28°57'15"N,  118°05'19°W,  est.  fishing  depth  0-600  m,  bot- 
tom depth  2,926-2,012  m,  IKMT,  VELERO  sta.  12494,  21  Nov. 
1968.  — AMNH  12902  (4,  55-70  HL,  250-305+  TL),  San  Cris- 
tobal Bay,  27°07'08"gN,  114°33'10"W,  519  m,  ALBATROSS  sta. 
5675.  15  Mar.  1911. 

Galapagos:  USNM  135605  ( 1,  77  HL.  346  TL),  ALBATROSS. 

Ecuador:  CAS  441%  (1,  14  HL,  67+  TL),  4°14'S,  81°26'W, 
0-1,830  m,  IKMT.  ANTON  BRUUN  cr.  18B,  sta.  756B  (field 
no.  LWK66-113).  8  Sep.  1966. 

Peru:  LACM  33883  (1,  66  HL.  303  TL),  6°42'S.  80°59'05"W. 
780m,  sta.  SNP1-25,  22  Jan.  1974. —LACM  33886(1.  41  HL. 
168  TL).  7°44'05"S,  80°30'05"W,  750-760  m,  sta.  SNP1-28,  23 
Jan.  1974.  —CAS  38324  (7,  48-66  HL.  180+  to  280  TL). 
7°49'S.  80°38'W.  605-735  m,  ANTON  BRUUN  cr.  18B.  sta.  754 
(field  no.  LWK66-93),  5  Sep.  1966.  —CAS  41197  (2,  13.5- 
14.7  HL,  66-73  TL),  11°53'S,  78°05'W,  0-1,125  m,  IKMT. 
ANTON  BRUUN  cr.  16,  sta.  656-0,  14  June  1%6.  — IMARPE 
uncat.  (1,  74  HL.  290+  TL),  18°10'S,  71°29'W,  610  m.  23  Aug. 
1972. 


Nezumia  pudens  Gilbert  and  Thompson 

(Figures  6b,  14c,  \Sh) 

Nezumia  pudens  GILBERT  AND  THOMPSON,  1916:472-473,  pi. 

5,  fig.  2  (types  from  ALBATROSS  sta.  2791  off  Lota,  Chile, 

38°08'S,  75°53'W,  1,240m). 
Lionurus  (Nezumia)  pudens:  GILBERT  AND  HUBBS  1916:146 

(name  only). 
Macruroplus  pu dens :  FOWLER  1944:48,  fig.  (list). 

DIAGNOSIS. — A  species  of  Nezumia  with  10- 
1 1  pelvic  fin  rays.  Gill-rakers  on  first  and  second 
arches  9-10  total.  Spinules  on  body  scales  slen- 
der, sharp,  conical,  in  dense  cluster  on  exposed 
fields.  Head  surface  fully  scaled  except  for  small 
area  along  ventromedian  line  of  snout  and  over 
gill  membranes.  Barbel  20-25  percent  of  HL; 
upper  jaw  30-39  percent  HL. 

DESCRIPTION. — General  features  offish  seen 
in  Figure  14a.  Head  rather  compressed,  surfaces 
smoothly  rounded.  Suborbital  ridge  low,  round- 
ed. Snout  short,  blunt  narrow;  blunt  scutes  at 
terminal  and  lateral  angles  not  especially  spiny 
or  set  off  from  adjacent  scales.  Mouth  subter- 
minal,  rather  short;  posterior  corners  somewhat 
restricted  by  lip  folds;  lips  thick,  papillaceous, 
especially  in  large  adults.  Barbel  rather  thick 
throughout,  not  tapering  to  filamentous  tip.  In- 
teropercle  broadly  exposed  posteriorly  and  ven- 
trally.  Posterodorsal  corner  of  opercle  angular, 
without  broad  tablike  extension.  Gill  mem- 
branes rather  broadly  connected  over  isthmus, 
with  a  narrow  free  fold.  Anus  located  in  middle 
third  of  distance  between  pelvic  fin  base  and 
anal  fin  origin.  Anterior  dermal  window  small, 
round,  lying  between  or  slightly  in  advance  of 
line  connecting  insertion  of  pelvic  fins.  Gill  fil- 
aments well  developed  but  relatively  short; 
length  of  longest  less  than  diameter  of  eye  lens, 
about  0.6  into  least  suborbital  width.  Pyloric 
caeca  slender,  numerous,  37,  39,  and  44  in  three 
specimens,  branched  two  or  three  times  at  base; 
length  of  longest  about  equal  to  interorbital 
width. 

Exposed  fields  on  body  scales  densely  cov- 
ered with  relatively  erect,  slender,  stiletto- 
shaped  spinules  arranged  in  quincunx  pattern  or 
in  close,  strongly  convergent  rows.  Almost  all 
of  head  uniformly  covered  with  small,  relatively 
adherent  scales  (Fig.  6b).  Small  area  along  ven- 
tral midline  and  ventral  margin  of  snout  naked. 
Scales  along  dorsal  half  of  suborbital  region 
heavier,  larger,  more  adherent,  and  in  two  dis- 
tinct rows.  Mandibles  and  exposed  surfaces  of 


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a 


FIGURE  14.  (a)  Nezumia  pudens  (Gilbert  and  Thompson).  CAS  38310.  50  mm  HL,  from  off  Chile  in  750  m.  (b)  Nezumia 
ventralis  Hubbs  and  Iwamoto,  new  species.  Holotype,  CAS-SU  68361,  26.4  mm  HL.  taken  off  the  Galapagos  in  549  m.  (r) 
Nezumia  orbitalis  (Carman),  LACM  33885-2,  48  mm  HL,  off  Peru  in  800  m. 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


165 


interopercle  covered  with  small  scales,  but  gill 
membranes  naked. 

Fins  moderate  in  size.  First  spinous  ray  of 
dorsal  fin  slender,  sharp,  spikelike;  second  spi- 
nous ray  slightly  prolonged,  armed  along  leading 
edge  with  slender,  sharp  spikes  spaced  suffi- 
ciently apart  so  that  spikes  do  not  overlap.  Outer 
ray  of  pelvic  fins  with  filamentous  tip  extending 
well  beyond  origin  of  anal  fin.  First  dorsal  fin 
placed  rather  far  posteriad;  origin  about  on  same 
vertical  as  anus  and  about  one  eye-lens  diameter 
behind  vertical  through  origin  of  pectoral  fins. 

Jaw  teeth  in  rather  broad,  coarse  bands  that 
narrow  posteriorly.  Outer  series  of  upper  jaw 
slightly  enlarged,  particularly  along  anterior  bor- 
der. 

Coloration  in  alcohol  medium  brown  with  ab- 
dominal and  cheek  area  blackish  tinged  with  vi- 
olet. Gill  membranes  black;  naked  margins  of 
snout,  nostrils,  lower  jaw,  opercular  bones,  and 
rim  of  orbit  blackish  or  darkly  swarthy.  Fins  all 
blackish;  distal  and  anterior  portions  of  mem- 
branes of  first  dorsal  fin  more  intensely  blackish. 
Lining  of  mouth  and  gullet  blackish;  that  of  gill 
cavity  blackish  along  external  margins  but  pale 
towards  inner  portions;  that  of  abdominal  cavity 
black  to  swarthy. 

COUNTS  (from  7  specimens). —  ID.  11,9-11; 
IP.  19-24;  2P.  10-11.  Medial  gill-rakers  on  first 
arch  1-2  +  7-9,  usually  2  +  8  (9-10  total);  on 
second  arch  1-2  +  8-9  (9-10  total).  Scale  rows 
(from  4  specimens)  below  first  dorsal  W/2  to  12; 
below  second  dorsal  8'/i  to  lO'/i;  below  midbase 
of  first  dorsal  ll/2  to  8!^>;  over  distance  equal  to 
predorsal  length  43^6. 

MEASUREMENTS  (from  6  specimens;  22-mm- 
HL  specimen  excluded). — Total  length  165^00 
mm;  head  length  32-68  mm.  The  following  in 
percent  of  head  length:  postrostral  length  of 
head  74-76;  snout  length  28-30;  preoral  length 
18-22;  internasal  width  19-23;  orbit  diameter 
31-35;  interorbital  width  22-23;  postorbital 
length  of  head  40^48;  distance  orbit  to  angle  of 
preopercle  37-43;  suborbital  width  15-16;  upper 
jaw  length  30-39;  length  barbel  20-25;  length 
first  gill-slit  14-18;  preanal  length  143-158; 
greatest  body  depth  73-81;  1D.-2D.  interspace 
34_43;  height  first  dorsal  fin  80-93;  length  pec- 
toral fin  56-64;  length  pelvic  fin  44-58. 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
zumia pudens  is  readily  distinguished  from  its 
most  closely  related  eastern  Pacific  congeners 
N.  stelgidolepis,  N.  pulchella,  and  N.  liolepis 
by:  (1)  ventral  surface  of  snout,  suborbital  re- 


gion, and  mandibular  rami  scaled  (vs.  mostly 
naked  in  these  other  species);  (2)  mucous  pores 
in  these  areas  not  prominent  (vs.  prominent); 

(3)  posterodorsal  angle  of  opercle  not  produced 
into  a  short  tab  or  flap  (vs.  a  distinct  flap  or  tab); 

(4)  gill  filaments  relatively  short,  length  less  than 
diameter  of  eye  lens  (greater  than  eye  lens  in 
others);  (5)  first  dorsal  fin  originating  more  pos- 
teriad than  others;  and  (6)  buccal  cavity  com- 
pletely blackish  (mostly  pale  in  stelgidolepis  and 
pulchella,  but  blackish  in  liolepis).  Its  relation- 
ship to  other  Pacific  species  of  Nezumia  is  un- 
certain and  probably  distant. 

Nezumia  pudens  bears  some  resemblance  in 
general  physiognomy  of  the  head  and  body  to 
N.  sclerorhynchus  (Valenciennes,  1838),  and  N. 
suilla  Marshall  and  Iwamoto,  1973,  from  the 
Atlantic,  but  the  higher  pelvic  fin  ray  count  and 
the  more  complete  squamation  on  the  ventral 
aspects  of  the  head  in  N.  pudens  are  distinguish- 
ing. 

SIZE. — To  at  least  68  mm  HL  and  400  mm  TL. 

DISTRIBUTION. — Known  only  off  central 
Chile  between  latitudes  32°S  and  38°S,  in  580- 
1,238  m. 

MATERIAL  EXAMINED  (7  specimens,  all  from  Chile). — 
USNM  uncat.  (formerly  CAS  38317)  (1,  39  mm  HL,  240  mm 
TL),  32°08.5'S,  71°43'W,  960  m,  ANTON  BRUUN  cr.  18A,  sta. 
703  (field  no.  LWK66-^7),  12  Aug.  1966.  —CAS  38315  (2,  22- 
39  HL,  106+  to  234  TL),  32°17'S,  71°39.5'W,  580  m,  ANTON 
BRUUN  cr.  18A,  sta.  702  (field  no.  LWK66-44),  1 1  Aug.  1966. 
—CAS  38310  (2.  50-68  HL,  267-400  TL),  34°06.5'S, 
72°18.5'W,  750  m,  ANTON  BRUUN  cr.  18A,  sta.  687  (field  no. 
LWK66-25),  5  Aug.  1966.  —USNM  76860  (holotype,  32  HL, 
170  TL)  and  CAS-SU  22727  (paratype,  34  HL,  165  TL), 
38°08'S,  75°53'W,  1,238  m,  ALBATROSS  sta.  2791,  14  Feb. 
1888. 


Nezumia  ventralis  Hubbs  and  Iwamoto, 
new  species 

(Figure  146) 

DIAGNOSIS. — A  Nezumia  with  15  pelvic  fin 
rays  and  24-27  pectoral  fin  rays.  Mandibular 
rami,  ventral  surfaces  of  snout,  and  most  of  ven- 
tral surfaces  of  suborbital  region  naked;  mucous 
pores  over  these  surfaces  large.  Scales  small, 
about  9Y2  below  origin  of  second  dorsal  fin;  spi- 
nules  on  body  scales  slender,  conical,  greatly 
reclined,  in  parallel  to  subparallel  rows. 

DESCRIPTION  OF  HOLOTYPE. — General  fea- 
tures offish  seen  in  Figure  14b.  Head  and  trunk 
moderately  compressed  and  deep.  Suborbital 
region  divided  into  an  upper  portion  covered 
with  two  longitudinal  rows  of  stout  scales  and 


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a  lower  portion  with  almost  no  scales;  the  sur- 
faces of  the  two  halves  meet  along  the  suborbital 
ridge  at  an  obtuse  angle.  Opercular  opening 
moderately  restricted,  extends  ventrally  to  be- 
low a  point  anteriad  of  preopercular  ridge.  Gill 
membranes  restricted,  without  a  free  fold  over 
isthmus.  Trunk  short,  distance  isthmus  to  anal 
fin  origin  less  than  postrostral  length  of  head. 
Periproct  moderate  in  size,  anus  much  closer  to 
insertion  of  pelvic  fins  than  to  origin  of  anal  fin. 
Ventral  light  organ  well  developed;  a  large,  oval 
dermal  window  forms  anterior  extension  of  peri- 
proct;  dermal  window  extends  forward  almost 
to  a  line  connecting  pelvic  fin  insertions.  Gill 
filaments  long,  well  developed;  longest  filament 
about  3.5-4.0  mm,  about  as  long  as  diameter  of 
eye  lens.  About  23  slender  pyloric  caeca. 

Teeth  in  moderately  broad  bands  in  both  jaws; 
bands  narrower  posteriorly  on  jaws.  Outer  se- 
ries of  teeth  in  upper  jaw  slightly  enlarged. 

Scales  of  body  covered  with  slender,  conical, 
reclined  spinules  arranged  in  parallel  to  subpar- 
allel  ridgelike  rows  that  give  a  somewhat  striated 
appearance  to  body  surfaces.  Four  to  five  rows 
of  spinules  on  larger  body  scales.  Spinules  on 
head  scales  more  erect  than  those  on  body 
scales,  and  spinules  often  arranged  in  divergent 
rows.  Scales  behind  bases  of  paired  fins,  behind 
first  dorsal  fin,  and  beneath  outer  margin  of  gill 
covers  without  spinules.  Scales  dorsally  on  sub- 
orbital  region  stout,  adherent,  and  covered  with 
coarse  spinules.  Scales  at  tip  and  lateral  angles 
of  snout  large,  tuberclelike,  and  coarsely  spined. 
Exposed  posterior  tip  of  interopercle  scaled. 
Mandibular  rami,  ventral  surfaces  of  snout,  and 
most  of  ventral  surface  of  suborbital  region  na- 
ked; mucous  pores  large  and  prominent  in  these 
areas.  There  appear  to  be  some  scale  pockets 
along  lowermost  branchiostegal  rays  of  holo- 
type,  but  condition  of  specimen  left  doubts  as 
to  their  actual  presence  (the  "pockets"  may 
simply  be  torn  skin). 

Fins  generally  well  developed.  First  dorsal  fin 
high,  length  almost  as  long  as  head  length  (long- 
est ray  broken  at  tip  in  holotype  and  paratype). 
Leading  edge  of  spinous  second  ray  armed  with 
sharp,  widely  spaced  denticles.  Outer  ray  of  pel- 
vic fin  slightly  prolonged,  extends  beyond  origin 
of  anal  fin.  Pelvic  and  anal  fins  originate  rela- 
tively far  forward;  pelvic  fin  origin  below  sub- 
opercle,  anal  fin  origin  below  hind  third  of  first 
dorsal  fin  base. 


Holotype  too  badly  faded  for  accurate  de- 
scription of  coloration.  See  color  description  of 
paratype. 

DESCRIPTION  OF  PARATYPE. — The  single 
paratype  is  in  poor  condition  and  falling  apart. 
The  snout  region  is  destroyed,  and  the  posterior 
half  of  the  tail  has  broken  off  and  almost  com- 
pletely disintegrated.  The  fins  and  scales  are  in- 
tact, however,  and  most  of  the  diagnostic  fea- 
tures of  the  species  could  be  examined.  The 
specimen  is  notably  larger  than  the  holotype 
(postrostral  length  of  head  31.0  mm  compared 
with  20.8  mm)  and  the  two  exhibit  some  onto- 
genetic  differences.  Thus,  the  spinules  on  scales 
are  more  numerous  than  in  the  holotype,  and 
the  larger  spinules  on  body  scales  tend  to  be 
lanceolate  rather  than  conical.  Spinule  rows  on 
body  scales  are  also  more  numerous,  six  to  eight 
on  the  larger  scales  of  paratype  compared  with 
four  to  five  in  comparable  scales  of  holotype. 

The  paratype  had  29  pyloric  caeca  compared 
with  the  approximate  count  of  23  obtained  in  the 
holotype  (I  did  not  enlarge  the  original  slit  in  the 
abdominal  wall  of  the  holotype  for  fear  of  dam- 
aging the  specimen  further).  Two  spinulated 
scales  were  present  at  the  base  of  the  left  fourth 
branchiostegal  ray,  but  none  were  on  the  op- 
posite member. 

Ground  color  of  paratype  dark  brown  with 
violet  tinge  over  abdomen.  Fin  membranes,  gill 
membranes,  and  naked  surfaces  of  snout,  sub- 
orbital  region,  and  mandibular  rami  chocolate- 
brown.  Gill  cavity  brown  on  inner  wall  and  along 
periphery  of  outer  wall  but  pallid  everywhere 
else.  Gill  filaments  and  gill  arches  pale,  the  tu- 
bercular rakers  dusky. 

COUNTS  (paratype  counts,  if  different,  in 
brackets).— ID.  11,10;  IP.  24 (left),  27 (right)  [25, 
25];  2P.  15.  Mesial  gill-rakers  on  first  arch  2  + 
9  [2  +  10],  on  second  arch  2  +  8  [2  +  9].  Scales 
below  first  dorsal  about  12  [about  13];  below 
second  dorsal  9l/2\  below  midbase  of  first  dorsal 
about  8  [about  9l/2]\  lateral  line  scales  over  dis- 
tance equal  to  predorsal  length  41  [not  taken  in 
paratype].  Pyloric  caeca  about  23  [29]. 

MEASUREMENTS  OF  HOLOTYPE. — Total  length 
142  mm,  head  length  26.4  mm.  The  following  in 
percent  of  head  length:  postrostral  length  of 
head  79.0;  snout  length  23.8;  preoral  length  20.4; 
internasal  width  19.7;  orbit  diameter  36.7;  inter- 
orbital  width  22.0;  postorbital  length  of  head 
38.2;  distance  orbit  to  angle  of  preopercle  34.0; 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


167 


suborbital  width  12.1;  upper  jaw  length  27.2; 
barbel  length  14.0;  length  first  (outer)  gill-slit 
15.9;  preanal  length  129;  isthmus  to  anus  47.4; 
outer  pelvic  ray  to  anal  origin  34.0;  greatest 
body  depth  68;  depth  over  anal  origin  57;  1D.- 
2D.  interspace  34;  length  pectoral  fin  about  53; 
length  outer  pelvic  ray  about  53. 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
zumia ventralis  appears  most  closely  related  to 
two  Atlantic  species,  N.  sclerorhynchus  (Valen- 
ciennes, 1838)  and  N.  suilla  Marshall  and  Iwa- 
moto,  1973.  The  combination  of  general  head 
and  body  shape,  the  presence  of  broad  naked 
areas  on  the  ventral  surfaces  of  the  snout  and 
suborbital  region,  the  naked  mandibular  rami, 
the  large  pores  on  the  head,  the  general  color- 
ation, and  the  placement  and  size  of  the  fins  sug- 
gest this  relationship.  The  new  species  is  readily 
distinguished  from  the  two  by  its  high  pelvic  fin 
ray  count  of  15,  compared  with  7  in  suilla  and 
usually  9  (rarely  7,  8,  or  10)  in  sclerorhynchus. 
N.  propinqua  (Gilbert  and  Cramer,  1897)  is  the 
only  other  species  of  the  genus  having  a  pelvic 
fin  ray  count  as  high  as  that  of  N.  ventralis,  but 
N.  ventralis  lacks  the  distinct  black  blotch  on 
the  tip  of  the  first  dorsal  fin  that  is  so  prominent 
in  propinqua,  and  it  has  more  pectoral  fin  rays 
(23  or  fewer  in  propinqua),  a  shallower  body,  a 
more  anteriorly  placed  pelvic  fin,  and  fewer 
scale  rows  below  the  second  dorsal  fin. 

DISTRIBUTION. — The  species  is  known  from 
only  two  specimens  taken  by  the  ALBATROSS  in 
the  Galapagos  Islands. 

ETYMOLOGY. — From  the  Latin  ventralis, 
adj.,  of  or  belonging  to  the  belly — in  reference 
to  the  ventral  light  organ. 

REMARKS. — The  paratype  was  first  examined 
(circa  1920)  by  Dr.  Carl  L.  Hubbs,  who  recog- 
nized the  distinctness  of  the  specimen  and  gave 
it  the  manuscript  name  of  "Lionurus  ventralis." 
Because  of  other  commitments,  Dr.  Hubbs  was 
unable  to  formally  describe  the  species  at  the 
time  and  consequently  returned  the  specimen  to 
the  National  Museum  along  with  other  macrou- 
rids  collected  by  the  ALBATROSS  in  the  eastern 
Pacific  in  1888  and  1904. 

MATERIAL  EXAMINED.— Holotype:  CAS-SU  68361,  26.4 
mm  HL,  142  mm  TL,  Galapagos  Islands,  off  Hood  (Espanola) 
I.,  1°30'30"S,  89°39'W,  549  m,  ALBATROSS  sta.  4642,  7  Nov. 
1904. 

Paratype:  USNM  135345,  1  badly  deteriorated  specimen, 


Galapagos  Islands,  00°29'S,  89°54'30"W,  717  m,  ALBATROSS 
sta.  2818,  15  Apr.  1888. 

Nezumia  orbitalis  (Garman) 
(Figure  14c) 

Macrurus  orbitalis  GARMAN,  1899:207-208,  pi.  47,  figs.  1-lb 
(original  description;  Gulf  of  Panama,  ALBATROSS  sta.  3385, 
in  523  m). 

Lionurus  (Nezumia)  orbitalis:  GILBERT  AND  HUBBS  1916:146 
(listed). 

orbitalis:  MAKUSHOK  1967:table  18. 


DIAGNOSIS.  —  A  species  of  Nezumia  with  9 
pelvic  fin  rays.  Gill-rakers  on  first  arch  1-2  +  8- 

10  (10-11  total);  on  second  arch  1-3  +  7-9  (9- 

11  total).  Spinules  on  body  scales  slender,  con- 
ical to  narrowly  lanceolate;  no  enlarged  middle 
row  of  spinules  on  scales.  Barbel  length  6-12 
percent  HL;  upper  jaw  length  27-32  percent 
HL.  First  dorsal  fin  with  distinct  black  tip. 

DESCRIPTION.  —  General  features  of  fish  seen 
in  Figure  14c.  A  petite  species  probably  not  ex- 
ceeding 200  mm  in  total  length.  Head  about  21- 
25  percent  of  total  length  in  specimens  with 
complete  tails.  Snout  pointed;  head  ridges  rela- 
tively well  developed  with  spinous  tubercles  at 
terminal  and  lateral  points  of  snout.  Dorsal  pro- 
file arched  high  over  nape  to  give  distinctive 
humpbacked  appearance.  Gill  membranes 
broadly  attached  across  isthmus,  with  only  a 
narrow  posterior  free  fold.  Anterior  dermal  win- 
dow of  light  organ  small,  circular,  situated  in  a 
shallow  fossa  between  bases  of  pelvic  fins;  anus 
close  behind. 

Spinous  second  ray  of  first  dorsal  fin  armed 
along  leading  edge  with  small,  closely  spaced 
teeth.  Outer  pelvic  ray  slightly  prolonged,  ex- 
tends past  origin  of  anal  fin.  Pelvic  fin  origin 
forward  of  pectoral  fin  origin,  which  in  turn  is 
slightly  forward  of  first  dorsal  fin  origin. 

Body  scales  with  slender,  conical  to  narrowly 
lanceolate  spinules  densely  covering  exposed 
fields,  disposed  in  as  many  as  15  rows  in  large 
specimens,  with  spinule  rows  converging  to- 
wards middle  of  field;  no  enlarged  middle  row 
of  spinules.  Most  ventral  surfaces  of  snout  and 
suborbital  region  naked.  Lower  jaw  surfaces  na- 
ked except  for  a  line  of  loose  scales  along  mid- 
line  of  each  ramus. 

Coloration.  Garman  (1899:208)  has  given  a 
good  color  description  from  the  types:  "Black- 
ish over  the  body  cavity;  reddish  brown  over 
the  muscular  portions  of  the  caudal  section; 
lighter  to  translucent  on  the  head  and  snout;  en- 


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tire  surface  tinted  with  grayish,  from  the  spi- 
nules  of  the  scales.  Fins  light;  first  dorsal  and 
pectorals  tipped  with  black.  Orbit  surrounded 
by  a  narrow  border  of  black."  The  peritoneal 
lining  of  the  lectotype  is  silvery  with  bold  punc- 
tations. 

Dentition  of  jaws  consists  of  small  teeth  in 
moderately  wide  bands  with  outer  series  of  up- 
per jaw  slightly  enlarged. 

COUNTS  (from  15  specimens). —  ID.  11,9-10 
(11  in  one  specimen);  IP.  20-22  (jf  =  20.8);  2P. 
9.  Gill-rakers  on  first  arch  1-2  +  8-10  (9-11  to- 
tal; x  =  10.3);  on  second  arch  1-3  +  7-9  (9-11 
total;  x  -  10.2).  Scales  below  first  dorsal  7-9; 
below  second  dorsal  6l/2  to  7;  below  midbase  of 
first  dorsal  4l/2  to  5l/i;  lateral-line  scales  over 
distance  equal  to  predorsal  length  of  head  36- 
45.  Pyloric  caeca  17,  17,  and  23  in  three  speci- 
mens. 

MEASUREMENTS  (from  15  specimens). — Total 
length  115-186  mm;  head  length  27-41  mm.  The 
following  in  percent  of  head  length  [range  (x\  n\ 
S.D.)]:  postrostral  length  of  head  73-78  (76.0; 
14;  1.47);  snout  length  25-30  (27.8;  15;  1.27); 
preoral  length  20-25  (22.9;  15;  1.81);  internasal 
width  18-22  (19.9;  15;  0.97);  orbit  diameter  29- 
34  (31.9;  15;  1.28);  interorbital  width  17-19 
(17.7;  15;  0.72);  postorbital  length  39-45  (41.5; 
13;  2.11);  orbit  to  angle  of  preopercle  33-40 
(34.5;  15;  1.96);  suborbital  width  11-14(12.4;  15; 
0.83);  upper  jaw  length  27-32  (28.5;  15;  1.55); 
length  barbel  6-12  (9.0;  13;  1.59);  length  outer 
gill-slit  14-18  (15.9;  15;  1.33);  preanal  length 
119-146  (123.8;  15;  12.42);  outer  pelvic  ray  to 
anal  origin  28-41  (33.9;  15;  5.19);  anus  to  anal 
origin  14-29(21.5;  15;  4.81);  greatest  body  depth 
63-82  (72.2;  15;  6.45);  1D.-2D.  interspace  34-57 
(44.1;  15;  7.35);  height  first  dorsal  fin  75-90 
(80.4;  10;  6.15);  length  pectoral  fin  48-78  (55.6; 
12;  7.35);  length  pelvic  fin  41-58  (49.2;  1 1;  4.49). 

RELATIONSHIPS  AND  COMPARISONS. — Ne- 
zumia  orbitalis  is  closely  related  to  N.  latiros- 
trata  and  the  two  have  narrow,  coincident,  geo- 
graphic ranges,  although  N.  orbitalis  is  usually 
found  at  shallower  levels  than  N.  latirostrata 
(the  two  species  were  represented  together  in 
only  one  haul,  SNP1-28,  off  Peru  in  800  m,  the 
deepest  record  for  N.  orbitalis).  Meristic  and 
morphological  features  of  the  two  species  are 
close,  and  the  general  appearance  is  similar. 
Color  differences  are  the  most  noticeable.  The 
first  dorsal  fin  of  N.  orbitalis  is  distinctly  pallid 
basally  and  black  tipped  distally,  whereas  in  N. 


latirostrata  the  fin  is  uniformly  blackish.  The 
head  of  A/,  orbitalis  is  paler  than  that  of  N.  la- 
tirostrata, and  the  prominent  black  orbital  ring 
of  the  former  is  essentially  lacking  in  the  latter. 
The  area  ventrally  on  the  naris  appears  smudged 
with  black  in  N.  latirostrata  but  is  scarcely  pig- 
mented  in  A/,  orbitalis.  Silvery  reflections  per- 
sist on  the  preopercle  of  A/,  orbitalis  specimens 
but  not  in  A/,  latirostrata  specimens,  and  the 
belly  denuded  of  scales  has  a  more  bluish  cast 
in  A/,  orbitalis.  Nezumia  latirostrata  is  generally 
a  much  darker  fish. 

The  low  pelvic  fin  ray  count,  the  number  of 
gill-rakers,  and  the  short  barbel  together  serve 
to  distinguish  A/,  orbitalis  from  other  eastern 
Pacific  members  of  the  genus. 

The  species  appears  close  to  the  widespread 
Atlantic  species  A/,  aequalis  (Giinther),  the  two 
sharing  many  meristic,  morphometric,  and  col- 
oration features.  Even  the  humpbacked  appear- 
ance of  A/,  orbitalis  is  shared  with  one  popula- 
tion of  A/,  aequalis — that  population  being 
coincidentally  also  one  from  the  eastern  side  of 
an  ocean  (the  Gulf  of  Guinea)  (see  Marshall  and 
Iwamoto  in  Marshall  1973:641).  A/,  orbitalis  is 
readily  distinguished  from  A/,  aequalis,  how- 
ever, by  its  somewhat  shorter  snout  length  (20- 
25  percent  HL  cf.  25-32  in  aequalis),  its  rela- 
tively longer  head  length  compared  with  the  to- 
tal length  (head  length  21-25  percent  of  total 
length,  cf.  15-18),  and  its  smaller  adult  size 
(probably  less  than  200  mm  TL,  as  judged  from 
the  175-mm  length  of  the  ripe  female  lectotype, 
cf.  more  than  270  mm  TL  in  aequalis  [Marshall 
and  Iwamoto  in  Marshall  1973:643]). 

DISTRIBUTION. — Gulf  of  Panama  to  Peru,  in 
523-800  m. 

SIZE.— To  at  least  41  mm  HL  and  186  TL. 

MATERIAL  EXAMINED  (15  specimens,  4  localities). — Pana- 
ma: MCZ  28578  (lectotype,  41  mm  HL,  175+  mm  TL),  MCZ 
50956  (5  paralectotypes,  formerly  MCZ  28578,  28-35  HL, 
115+  to  153  TL),  and  USNM  57856  (2  paralectotypes,  28-34 
HL,  129+  to  130+  TL),  all  from  Gulf  of  Panama,  7°32'36"N, 
79°16'W,  286  fms  (523  m),  ALBATROSS  sta.  3385,  8  Mar.  1891. 

Peru:  LACM  33885  (2,  40-40  HL,  174+  to  186  TL),  7°46'S, 
80°31'W,  800  m,  sta.  SNP1-27,  23  Jan.  1974.  —LACM  33886 
(4,  27-39  HL,  118-178  TL),  7°44'05"S,  80°30'05"W,  750-760 
m,  sta.  SNP1-28,  23  Jan.  1974.  — IMARPE  (1,  40  HL,  175  + 
TL),  13°57.2'S,  76°42.0'W,  580-600  m,  7  Aug.  1972. 

Nezumia  latirostrata  (Garman) 

(Figures  4b,  \5a,  \8e ) 

Macrurus  latirostratus  GARMAN,  1899:211-212,  pi.  48,  fig.  2 
(original  description;  types  from  ALBATROSS  sta.  3354,  3384 
and  3394  in  Gulf  of  Panama,  322-511  fms  [589-935  m]). 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


169 


a 


FIGURE  15.  (a)  Nezumia  latirostrata  (Garman),  CAS  38330,  40  mm  HL,  from  off  Ecuador  in  700-1,000  m.  (b)  Nezumia 
convergent  (Garman),  CAS  36816,  44  mm  HL,  from  off  Ecuador  in  850-1 .400  m.  (c)  Nezumia  loricata  atomos  new  subspecies. 
Holotype,  CAS  38312,  51  mm  HL,  297+  TL,  from  off  Chile  in  1,170-1,480  m. 


Lionurus   (Nezumia)   latirostratus:   GILBERT  AND  HUBBS 

1916: 146  (name  only). 
Nezumia  latirostrata:  MAKUSHOK  1967:table  18. 

DIAGNOSIS. — A  species  of  Nezumia  with  9- 
10  pelvic  fin  rays.  Gill-rakers  on  first  and  second 
arch  9-11  total  (7-9  on  lower  limb).  Barbel 
length  5-10  percent  of  HL;  upper  jaw  length  27- 


32.  Spinules  on  body  scales  lanceolate,  arranged 
in  indistinct  rows  without  enlarged  middle  row. 
Scales  below  first  dorsal  1V2  to  9;  below  second 
dorsal  6  to  1V2.  First  dorsal  fin  uniformly  black- 
ish. 

DESCRIPTION. — General  features  of  fish  seen 
in  Figure  15a.  Head  and  trunk  moderately  com- 


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pressed,  tail  greatly  so.  Head  ridges  prominent, 
suborbital  ridge  strongly  angular.  Mouth  subter- 
minal;  rictus  extends  to  below  anterior  edge  of 
eye  lens;  maxilla  extends  to  below  middle  of 
eye.  Terminal  and  lateral  angles  of  snout  armed 
with  small  but  stout,  spiny  tubercular  scales.  In- 
terorbital  region  slightly  concave,  width  about 
60  percent  of  orbit  diameter.  Gill  membranes 
broadly  attached  over  isthmus  (Fig.  4b),  with  a 
narrow  posterior  free  fold;  gill  opening  extends 
anteroventrally  slightly  anteriad  of  a  vertical 
through  posterior  edge  of  preopercle.  Body 
moderately  deep  in  adults,  generally  more  slen- 
der in  the  young;  greatest  body  depth  more  than 
postrostral  length  of  head  in  all  specimens  larger 
than  about  30  mm  HL.  Light  organ  externally 
represented  by  a  small,  black,  oblate,  naked  fos- 
sa between  bases  of  pelvic  fins  and  anterior  to 
anus.  Anus  situated  notably  closer  to  insertion 
of  pelvic  fins  than  to  origin  of  anal  fin.  Gill  fil- 
aments short;  length  of  longest  slightly  shorter 
than  diameter  of  eye  lens. 

First  dorsal  fin  of  moderate  height,  length  of 
longest  ray  less  than  head  length;  second  spi- 
nous  ray  slightly  produced  into  a  thin  filament; 
leading  edge  of  ray  armed  with  small,  closely 
spaced  teeth;  in  some  specimens,  tip  of  each 
tooth  overlaps  base  of  adjacent  tooth.  Paired 
fins  of  moderate  size;  outer  ray  of  pelvic  fin  pro- 
duced, extends  posteriad  well  beyond  origin  of 
anal  fin. 

Body  scales  moderately  deciduous,  as  judged 
from  generally  denuded  state  of  specimens  ex- 
amined. Spinules  on  body  scales  distinctly  lan- 
ceolate, densely  covering  exposed  fields,  ar- 
ranged in  indistinct  longitudinal  rows  that 
converge  towards  middle  of  field.  Scales  absent 
on  ventral  surface  of  snout,  but  surface  copi- 
ously dotted  with  dark  sensory  papillae.  Sub- 
orbital  region  below  ridge  with  a  wedge-shaped 
cluster  of  small  scales.  Mandibles  with  small 
loose  scales  along  midline  of  each  ramus  but 
naked  at  anterior  end. 

Teeth  small,  in  bands  of  moderate  width  in 
both  jaws.  Outer  series  of  teeth  in  upper  jaw 
scarcely  enlarged. 

Coloration  in  alcohol  swarthy.  Bluish  tinge 
over  trunk  and  violet  tinge  over  head  and  tail 
pronounced  in  denuded  specimens.  All  fins  dark 
brown  to  brownish  black.  Lining  of  oral  cavity 
dusky,  that  of  gill  cavities  black  except  for  pale 
areas  ventrally  and  along  medial  wall.  Peritoneal 


cavity  pale  but  peppered  with  large  melano- 
phores. 

COUNTS  (from  57  specimens). —  ID.  11,9-10 
(11,8  in  two  specimens,  11,11  in  two  others);  IP. 
18-23  (x  =  20.65;  n  =  105;  S.D.  =  1.0);  2P.  9- 
10.  Gill-rakers  on  first  and  second  arch  1-3  +  7- 
9  (total  9-11).  Scales  below  first  dorsal  7!/2  to  9 
(rarely  7);  below  second  dorsal  6  to  7l/2\  lateral- 
line  scales  over  distance  equal  to  predorsal 
length  of  head  32-38  (jt  =  35.88;  n  =  34;  S.D.  = 
1.5).  Pyloric  caeca  17-25  (x  =  21.23;  n  =  30; 
S.D.  =  2.4). 

MEASUREMENTS  (from  56  specimens). — Total 
length  128-231  mm;  head  length  26^8  mm.  The 
following  in  percent  of  head  length  [range  (x; 
n;  S.D.)]:  postrostral  length  of  head  71-77  (73.53; 
56;  1.78);  snout  length  27-34  (29.97;  56;  4.36); 
preoral  length  23-31  (26.16;  56;  1.78);  internasal 
width  20-26  (23.39;  55;  1.23);  orbit  diameter  30- 
36  (33.78;  55;  1.46);  interorbital  width  17-25 
(20.84;  56;  1.57);  postorbital  length  of  head  36- 
43  (38.94;  54;  1.52);  orbit  to  angle  of  preopercle 
33-39  (35.25;  55;  1.36);  suborbital  width  13-17 
(15.18;  56;  1.11);  upper  jaw  length  27-32  (29.55; 
56;  1.30);  length  barbel  5-10  (8.19;  49;  1.57); 
length  first  gill-slit  11-16  (13.59;  54;  1.13);  pre- 
anal  length  123-157  (140.93;  56;  7.57);  outer  pel- 
vic ray  to  anal  fin  origin  27-48  (37.14;  55;  4.59); 
anus  to  anal  fin  origin  16-33  (22.27;  52;  3.78); 
greatest  body  depth  66-96  (77.22;  56;  11.01); 
1D.-2D.  interspace  34-55  (one  specimen  with 
72)  (45.41;  56;  6.30);  height  first  dorsal  fin  75-98 
(83.78;  18;  6.10);  length  pectoral  fin  52-68 
(59.32;  50;  4.96);  length  pelvic  fin  41-67  (55.57; 
49;  6.92). 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
zumia  latirostrata  and  N.  convergens  are 
closely  related  and  specimens  of  the  two  are  dif- 
ficult to  differentiate.  Chief  differences  between 
the  two  species  lie  in  (1)  gill-raker  counts,  (2) 
barbel  length,  (3)  scales  below  first  dorsal,  (4) 
scales  over  distance  equal  to  predorsal  length, 
and  (5)  body  length.  Nezumia  latirostrata  gen- 
erally has  7-9  gill-rakers  on  the  lower  limb  of 
the  first  and  second  arches,  whereas  N.  conver- 
gens has  5-7.  Barbel  lengths  contrast  at  5-10 
percent  HL  for  N.  latirostrata  and  8-20  percent 
(12-18  in  most)  for  N.  convergens.  Scale  rows 
below  the  origin  of  the  first  dorsal  fin  are  less 
numerous  in  latirostrata  (7l/2  to  9)  than  in  con- 
vergens (8'/2  to  10,  a  few  individuals  with  11). 
The  number  of  anterior  lateral-line  scales  over 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


171 


a  distance  equal  to  the  predorsal  length  is  gen- 
erally lower  in  latirostrata  (32-38  in  most  spec- 
imens compared  with  36-42  in  convergens),  but 
there  is  considerable  overlap  with  convergens 
in  this  feature.  The  body  is  somewhat  shorter  in 
latirostrata;  the  ratio  HL:TL  is  between  1:4.5 
and  1:5.5  (fewer  than  12  percent  of  the  speci- 
mens with  an  intact  tail  had  a  ratio  of  more  than 
1:5.2),  whereas  in  convergens  it  is  1:5.3  to  1:5.8. 

Nezumia  latirostrata  also  closely  resembles 
N.  loricata.  The  principal  differences  between 
the  two  rest  in  the  pelvic  fin  ray  counts  (9-10  in 
latirostrata,  10-12  in  loricata);  barbel  length  (5- 
10  percent  HL  vs.  10-17);  number  of  scale  rows 
below  midbase  of  first  dorsal  fin  (usually  4l/2  to 
5Vi  vs.  6l/2  to  7Vi)  and  below  the  origin  of  the 
second  dorsal  fin  (6  to  ll/2  vs.  8-9);  and  the  ratio 
of  head  length  to  body  length  (1:4.8  to  1:5.5  vs. 
1:5.1  to  1:6.1). 

Nezumia  latirostrata  is  readily  distinguished 
from  other  eastern  Pacific  species  of  Nezumia 
by  characters  given  in  the  key.  Nezumia  lati- 
rostrata is  distinguishable  from  the  Hawaiian 
species  of  the  genus  by  a  combination  of  char- 
acters that  includes  snout  shape  (blunter  and 
shorter  in  holocentrus  (Gilbert  and  Cramer, 
1897);  burragei  (Gilbert,  1905);  hebetata  (Gil- 
bert, 1905);  and  ectenes  (Gilbert  and  Cramer, 
1897))  and  pelvic  fin  ray  count  (more  in  propin- 
qua  (Gilbert  and  Cramer,  1897)  and  obliquata 
(Gilbert.  1905)).  Of  the  Japanese  species  report- 
ed by  Okamura  (1970a),  all  but  four  (proxima 
(Smith  and  Radcliffe,  1912,  in  Radcliffe),  tomi- 
yamai  (Okamura,  1963),  burragei  (Gilbert, 
1905),  and  dara  (Gilbert  and  Hubbs,  1916))  can 
be  differentiated  from  latirostrata  by  the  num- 
ber of  pelvic  fin  rays,  but  N.  tomiyamai,  N.  bur- 
ragei, and  N.  dara  have  more  scale  rows  below 
the  origin  of  the  second  dorsal  fin  than  does  N. 
latirostrata,  and  N.  proxima  has  a  longer  barbel 
(less  than  two  times  into  orbit  diameter  com- 
pared with  three  or  more  times  in  latirostrata). 

Most  Atlantic  members  of  the  genus  (see  Mar- 
shall and  Iwamoto  in  Marshall  1973:624-649) 
are  distinguishable  from  N.  latirostrata  by  a 
combination  of  pelvic  fin  ray  count,  barbel 
length,  and  shape  of  scale  spinules.  The  western 
Atlantic  species  N.  cyrano  Marshall  and  Iwa- 
moto. 1973,  is  "probably  most  closely  related  to 
N.  latirostratus"  (ibid.:629),  but  the  two  differ 
in  snout  length  (longer  in  cyrano,  32-36  percent 
HL,  as  opposed  to  27-34,  most  specimens  below 


33,  in  latirostrata),  upper  jaw  length  (24-28  in 
cyrano  vs.  27-32  in  latirostrata),  and  length  of 
first  gill-slit  (6-10  vs.  11-16). 

DISTRIBUTION. — Panama  (SE  of  Punta  Mala) 
to  northern  Peru  (SE  of  Lobos  de  Afuera)  in  605- 
1,400  m. 

SIZE. — To  at  least  48  mm  HL  and  more  than 
234  mm  TL. 

MATERIAL  EXAMINED  (182  specimens  from  1 1  localities). — 
Panama:  USNM  57855  (2  syntypes,  39^1  mm  HL.  198+  to 
205  mm  TL),  7021'N,  79°35'W,  935  m,  ALBATROSS  sta.  3394, 
10  Mar.  1891.  — GCRL  14259  (2.  47-^8  HL.  215-198+  TL), 
7°18'N,  79°38'W,  732-805  m,  CANOPUS  sta.  1285,  24  Mar. 
1974.  —GCRL  14260(1,41  HL,  167+  TL),  7°13'N,  79°18'W, 
805-841  m,  CANOPUS  sta.  1291,  12  Mar.  1974.  — CAS-SU 
25246  (2,  44^*6  HL,  195+  to  245  TL).  "between  Galapagos 
and  Panama  ALBATROSS  1888"  (no  other  data). 

Ecuador:  CAS-SU  24091  (8,  28-41  HL)  and  CAS-SU  25223 
(9,  38-42  HL),  00°37'S,  81°00'W,  733  m.  ALBATROSS  sta.  2792, 
2  Mar.  1888.  —CAS  38328  (1,  42  HL,  175+  TL),  2°20'S, 
81°16'W.  850-1.400  m,  TE  VEGA  cr.  19.  sta.  84.  4  Aug.  1968. 
—CAS  38330  (49.  25^t5  HL,  133-215  TL).  2°25'S.  81°10'W. 
700-1,000  m.  TE  VEGA  cr.  19,  sta.  148,  1  Sep.  1968.  —CAS 
40826  (60.  38-45  HL,  190-234  TL),  3°15'S,  80°55'W,  945-960 
m,  ANTON  BRUUN  cr.  18B.  sta.  770  (LWK66-120),  10  Sep. 
1966. 

Peru:  USNM  1 18019(1.  42  HL.  215+ TL),  5°47'S,  81°24'W, 
1.030  m.  ALBATROSS  sta.  4653,  12  Nov.  1904.  — LACM  33885 
(2,  37-38  HL,  175-192  TL),  7°46'S,  80°31'W,  800  m.  sta. 
SNP1-27,  23  Jan.  1974.  —CAS  38321  (35,  16-46  HL,  81-205 
TL)  and  CAS  38322  (10,  27^6  HL,  100+  to  203  TL),  7°49'S. 
80°38'W,  605-735  m.  ANTON  BRUUN  cr.  18B.  sta.  754 
(LWK66-93),  5  Sep.  1966. 

In  addition,  the  following  ALBATROSS  specimens  were  ex- 
amined but  no  measurements  or  counts  were  taken  from  them: 
MCZ  28570  (6  syntypes),  sta.  3354;  MCZ  28571  (10  syntypes), 
sta.  3384;  and  MCZ  28572  (3  syntypes).  sta.  3394. 

Nezumia  convergens  (Garman) 

(Figures  156,  18/) 

Macrurus  convergens  GARMAN,  1899:210-211.  pi.  48,  fig.  1 
(Gulf  of  Panama.  695-1,020  fms  [1,271-1.865  m],  ALBA- 
TROSS sta.  3353,  3357.  and  3393). 

Macrurus  cuspidatus  GARMAN,  1899:209-210  (Gulf  of  Cali- 
fornia. 27°34'N,  110°53'40"W.  905  fms  [1.655  m].  ALBA- 
TROSS sta.  3436). 

Macrurus  trichiurus  GARMAN,  1899:215  (Gulf  of  Panama,  555 
fms  [1,014  m],  ALBATROSS  sta.  3358). 

Lionurus  (Nezumia)  convergens:  GILBERT  AND  HUBBS 
1916: 146  (listed). 

Lionurus  (Nezumia)  cuspidatus:  GILBERT  AND  HUBBS 
1916: 146  (listed). 

Lionurus  (Nezumia)  trichiurus:  GILBERT  AND  HUBBS  1916:146 
(listed). 

Nezumia  convergens:  MAKUSHOK  1967:table  18. 

Nezumia  cuspidata:  MAKUSHOK  l%7:table  18. 

Nezumia  trichiura:  MAKUSHOK  1967:table  18. 

Sphagemacrurus  trichiurus:  MARSHALL  1973:623  (listed). 

DIAGNOSIS. — A  species  of  Nezumia  with  pel- 


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vie  fin  rays  10-11.  Gill-rakers  on  first  arch  7-9 
(5-7  on  lower  limb);  on  second  arch  8-9.  Spi- 
nules  on  body  scales  slender,  conical  to  narrow- 
ly lanceolate,  in  discrete  longitudinal  rows,  mid- 
dle row  often  enlarged.  Barbel  length  8-20 
percent  HL  (usually  2.5-3.0  in  orbit);  upper  jaw 
length  26-32.  First  dorsal  fin  uniformly  blackish. 

DESCRIPTION. — General  features  offish  seen 
in  Figure  15b.  Head  slightly  compressed,  ridges 
of  head  generally  well  developed;  suborbital 
ridge  prominent.  Mouth  subterminal,  upper  jaw 
extends  posteriad  to  below  middle  of  eye.  Snout 
moderately  pointed;  terminal  and  lateral  angles 
with  stout  tubercles  armed  with  radiating  rows 
of  small  spinules.  Interorbital  region  flat  to 
slightly  concave,  narrow,  somewhat  more  than 
half  orbit  diameter.  Gill  membranes  broadly  at- 
tached over  isthmus,  with  only  a  narrow  pos- 
terior free  fold;  gill  openings  extend  anteroven- 
trally  to  beneath  posteroventral  angle  of 
preopercular  ridge  (Fig.  4b).  Body  slender; 
greatest  body  depth  (at  origin  of  first  dorsal  fin) 
less  than  postrostral  length  of  head;  7-8  in  total 
length  in  large  adults.  Light  organ  not  well  de- 
veloped externally;  no  scaleless  fossa  between 
pelvic  fins.  Anus  within  middle  third  of  distance 
between  pelvic  fin  insertion  and  anal  fin  origin. 
Gill  filaments  rather  short,  length  of  longest 
shorter  than  diameter  of  eye  lens. 

Second  spinous  ray  of  first  dorsal  fin  longer 
than  postrostral  length  of  head;  leading  edge 
armed  with  rather  closely  spaced,  nonoverlap- 
ping  slender,  sharp  teeth.  Pectoral  and  pelvic 
fins  originate  about  on  same  vertical,  which  is 
anterior  to  origin  of  first  dorsal  fin.  Outer  pelvic 
ray  in  adults  extends  slightly  past  anal  fin  origin. 
Second  dorsal  fin  rudimentary  throughout. 

Body  scales  somewhat  deciduous.  Slender, 
sharp,  conical  spinules  uniformly  cover  exposed 
fields,  aligned  in  slightly  convergent  longitudinal 
rows;  spinules  in  middle  row  usually  larger  than 
those  of  lateral  rows;  10-12  rows  of  spinules  on 
scales  of  large  adults.  Almost  all  of  ventral  snout 
surface  and  part  of  anteroventral  portion  of  sub- 
orbital  region  without  scales.  Mandibular  rami 
covered  posteriorly  with  narrow  band  of  small, 
loose  scales;  mandibles  generally  naked  ante- 
riorly, but  liberally  pocketed  there  with  small 
pores. 

Dentition  composed  of  small  teeth  in  broad 
bands  in  both  jaws;  outer  series  of  teeth  in  upper 
jaw  not  notably  enlarged. 

Coloration  in  alcohol  generally  brownish 


black  with  hint  of  violet  (especially  on  tail); 
bluish  tinge  over  abdomen.  Fins  uniformly  black 
or  brownish  black.  Oral  cavity  dusky;  peritoneal 
cavity  flesh  colored,  often  lightly  peppered  with 
large  melanophores. 

COUNTS  (from  80  specimens).  —ID.  11,9-10 
(rarely  8  or  11);  IP.  18-22  (x  =  20.18;  n  =  142; 
S.D.  =  1.02);  2P.  10-11  (rarely  9).  Gill-rakers 
on  first  and  second  arch  1-2  +  5-7  (total  7-9; 
.v  =  8.04;  n  =  80;  S.D.  =  0.56).  Scales  below 
first  dorsal  81^-10  (rarely  11);  below  second  dor- 
sal 5l/2-9  (usually  6'/2-9);  lateral  line  scales  over 
distance  equal  to  predorsal  length  of  head  33-44 
(usually  36-42).  Pyloric  caeca  21-32  (usually  25- 
30)  (jf  =  26.97;  n  =  35;  S.D.  2.55). 

MEASUREMENTS  (from  80  specimens,  25  lo- 
calities, 17.5-51.6  mm  HL,  97-300  mm  TL),  ex- 
pressed as  percentage  of  head  length  [range  (x; 
n;  S.D.)]:  postrostral  length  of  head  69-76 
(72.60;  79;  1.43);  snout  length  27-34  (30.46;  80; 
1.37);  preoral  length  22-31  (26.99;  78;  1.59);  in- 
ternasal  width  17-23  (20.50;  70;  1.31);  orbit  di- 
ameter 31-37  (33.74;  80;  1.31);  interorbital  width 
16-23  (19.73;  79;  1.29);  postorbital  length  34-40 
(37.78;  78;  1.59);  orbit  to  angle  of  preopercle  32- 
40  (35.17;  77;  1.72);  suborbital  width  12-17 
(14.41;  77;  0.96);  upper  jaw  length  26-32  (28.83; 
78;  7.37);  length  barbel  8-20  (14.08;  74;  2.64); 
length  first  gill-slit  11-19  (13.22;  76;  1.36);  pre- 
anal  length  133-181  (150.13;  73;  8.47);  distance 
outer  pelvic  ray  to  anal  origin  27-55  (40.11;  75; 
5.24);  vent  to  anal  origin  14-28  (20.54;  66;  3. 1 1); 
greatest  body  depth  60-78  (71 .21;  64;  5.23);  1D.- 
2D.  interspace  28-68  (45.74;  75;  9.00);  height 
first  dorsal  fin  68-93  (79.68;  53;  5.13);  length 
pectoral  fin  46-60  (52.30;  64;  3.52);  length  pelvic 
fin  36-61  (46.03;  59;  4.85). 

RELATIONSHIPS  AND  COMPARISONS. — In  the 
eastern  Pacific.  Nezumia  convergens  appears 
closest  to  N.  latirostrata  (Garman),  N.  loricata 
(Garman),  and  N.  orbitalis  (Garman),  but  it  is 
readily  distinguished  from  these  three  by  its  few- 
er gill-rakers  on  the  first  arch  (9  or  fewer  total, 
5-6  [rarely  7]  on  lower  limb,  as  compared  with 
9  or  more  total,  7-11  on  lower  limb).  Spinules 
on  body  scales  are  generally  more  slender  and 
conical  than  those  of  N.  latirostrata  and  N.  lor- 
icata, and  they  are  arranged  in  discrete  parallel 
rows  with  the  middle  row  often  enlarged  (this 
compares  with  spinules  in  a  somewhat  quincunx 
arrangement  or  in  rows  that  converge  strongly 
toward  the  midline  in  the  other  two  species).  In 
addition,  the  body  is  somewhat  more  slender 


IWAMOTO:   EASTERN  PACIFIC  MACROURIDAE 


173 


than  that  of  the  other  three  species  (greatest 
depth  less  than  75  percent  of  head  length  in  most 
adults  of  N.  convergens  compared  with  more 
than  75  percent  in  most  adults  of  other  species), 
the  anus  is  usually  more  posteriorly  placed, 
often  closer  to  the  anal  fin  origin  than  to  the 
insertion  of  the  pelvic  fins  (always  closer  to  the 
pelvic  fin  insertion  in  the  others),  and  the  ante- 
rior dermal  window  of  the  light  organ  is  poorly 
defined  externally  (well  defined  in  others).  The 
barbel  is  slightly  longer  than  that  of  N.  latiros- 
trata,  and  the  first  dorsal  fin  is  uniformly  black- 
ish, not  black  blotched  as  in  N.  orbitalis. 

Nezumia  convergens  seems  closely  related  to 
the  western  Atlantic  species  N.  suilla  Marshall 
and  Iwamoto,  1973,  with  which  it  shares  a  close 
similarity  in  general  physiognomy,  coloration, 
scale  spinulation,  and  extent  of  naked  areas  on 
the  ventral  surfaces  of  the  head.  The  lower  pel- 
vic fin  ray  count  (7)  and  broader  scale  spinules 
in  that  species,  nonetheless  readily  distinguish 
it  from  N.  convergens. 

REMARKS  ON  SYNONYMY. — In  my  studies  of 
the  grenadiers  described  by  Carman  (1899),  I 
have  encountered  three  instances  where  he  de- 
scribed different  specimens  of  a  single  species- 
group  taxon  as  different  species.  In  the  present 
instance,  the  nominal  species  concerned  are 
Macrurus  cuspidatus,  M.  convergens,  and  M. 
trichiurus.  As  first  revisor,  I  have  chosen  to  es- 
tablish the  name  M.  convergens  over  the  other 
two,  primarily  because  the  type-series  for  the 
species  is  large  and  representative,  while  M. 
cuspidatus  and  M.  trichiurus  were  each  de- 
scribed from  single  specimens,  the  last  from  a 
juvenile. 

The  holotype  of  M.  cuspidatus  agrees  well 
with  the  many  well-preserved  type-specimens  of 
M.  convergens  except  for  its  much  deeper  body. 
The  entire  trunk  area  of  the  holotype.  in  fact, 
appears  to  be  abnormally  swollen,  as  if  it  had 
been  over-injected  with  preservative.  If  this  is 
true,  the  deeper  body  is  of  no  consequence.  The 
Gulf  of  California  capture  of  M.  cuspidatus  rep- 
resents the  northernmost  record  of  the  species 
and  the  only  record  from  the  Gulf. 

The  holotype  of  M.  trichiurus  is  a  young  spec- 
imen (17.5  mm  HL,  111  mm  TL)  slightly  dam- 
aged ventrally.  It  agrees  in  all  salient  features 
with  small  specimens  of  N.  convergens  I  have 
examined.  Although  Carman  (1899:215)  gave  a 
pelvic  fin  ray  count  of  8  for  the  species,  I  count- 
ed 10  on  both  fins  of  the  holotype.  The  gill-raker 


count  on  that  specimen  was  1  +  6  on  the  mesial 
side  of  both  the  first  and  the  second  arches. 

DISTRIBUTION. — Nezumia  convergens  is  one 
of  the  most  numerous  and  widely  distributed 
grenadier  in  the  eastern  Pacific  Ocean.  The 
species  has  the  greatest  latitudinal  range  of  any 
Nezumia  in  the  eastern  Pacific  except  N.  stel- 
gidolepis,  although  present  collections  show  a 
broad  gap  between  the  Islas  Tres  Marias  and 
Costa  Rica.  This  gap  is  of  little  significance, 
however,  in  that  deep  trawling  along  the  Central 
American  coast  between  the  Gulf  of  Panama  and 
the  Gulf  of  California  has  been  extremely  lim- 
ited. One  Gulf  of  California  record;  others  from 
off  northern  Mexico.  Costa  Rica,  Panama,  Co- 
cos  Ridge,  Galapagos,  Ecuador,  Peru,  and  Chile 
(to  34°53.5'S).  Capture  depths  from  600  m  to 
1,865  m. 

SIZE.— To  at  least  50  mm  HL  and  300  mm  TL. 

MATERIAL  EXAMINED.— Mexico:  MCZ  28575  (1,  52  mm 
HL;  holotype  of  Macrurus  cuspidatus  Carman,  1899).  Gulf  of 
California  off  Guaymas,  Sonora,  27°34'N,  110°53'40"W.  1,655 
m,  ALBATROSS  sta.  3436,  22  Apr.  1891.  —  LACM  3 1124^  (3, 
43^t5  HL,  250-270  TL).  N  of  Islas  Tres  Marias,  21°52'30"N, 
106°47'36"W,  800-550  m,  VELERO  IV  sta.  13770.  21  Jan.  1970. 

Costa  Rica:  LACM  33588  (4,  28-50  HL,  161-295  TL), 
9°45'18"N,  85°52'24"W,  1,865-1,372  m,  VELERO  IV  sta.  18932, 
12-13  May  1973. 

Panama:  MCZ  28574  (1  damaged  syntype).  7°I5'N, 
79°36'W,  1,865  m,  ALBATROSS  sta.  3393,  10  Mar.  1891.  - 
GCRL  14258(1.  26  HL,  132+  TL),  7°13'N,  79°18'W,  805-841 
m,  CANOPUS  sta.  1291,  12  Mar.  1974.  —MCZ  28573  (3  syn- 
types,  31^2  HL,  170+  to  220+  TL),  7°06'15"N,  80°34'W, 
1,271  m,  ALBATROSS  sta.  3353,  23  Feb.  1891.  —MCZ  28556 
(1.  17.5  HL,  111  TL;  holotype  of  Macrurus  trichiurus  (Gar- 
man,  1899),  6°30'N,  81°44'W,  1,006  m,  ALBATROSS  sta.  3358, 
24  Feb.  1891. 

Cocos  Island:  AMNH  3471  (3,  29^*3  HL),  96  km  s  of  Cocos, 
ARCTURUS  sta.  74,  May  1925. 

Galapagos:  CAS  36814  (1,  47  HL,  248  TL),  27  km  SSE  of 
Isla  San  Cristobal,  1°06'S,  89°22'W.  700-800  m.  TE  VEGA  cr. 
19,  sta.  102,  12  Aug.  1968. 

Ecuador:  CAS  36817  (1.  47  HL,  265  TL),  off  Gulf  of  Gua- 
yaquil, 2°10'S,  STO'W,  800-1,000  m,  TE  VEGA  cr.  19,  sta. 
144,  31  Aug.  1968.  —CAS  36816  (8,  19-44  HL,  101-252  TL), 
2°20'S,  81°16'W,  850-1,400  m,  TE  VEGA  cr.  19.  sta.  84,  4  Aug. 
1968.  —CAS  36815  (2.  22-30  HL,  108-175  TL).  2°25'S, 
STIO'W,  700-1,000  m,  TE  VEGA  cr.  19.  sta.  148.  1  Sep.  1968. 
—CAS  36812  (35,  23^8  HL.  134-230+  TL),  3°15'S,  80°55'W, 
ANTON  BRUUN  cr.  18B,  sta.  770  (field  no.  LWK66-120). 

Peru:  CAS  36813  (3,  40-43  HL.  205-230  TL).  4°10'S, 
81°27'W,  1,815-1,860  m,  ANTON  BRUUN  cr.  18B,  sta.  766 
(field  no.  LWK66-115),9Sep.  1966. —CAS  28760  (1,  21  HL, 
97  TL),  17°08'05"S,  73°28'04"W,  860  m,  27  Jan.  1972.  — IM- 
ARPE  (1,  37  HL,  206+  TL),  18°17.3'S,  71°11'W,  600  m.  28 
Jan.  1972.— CAS  28762  (1,33  HL,  177  TL),  18°19'S,  71°12'W, 
810  m,  28  Jan.  1972.  — IMARPE  (1.  32  HL)  (bottle  label  illeg- 
ible), 800  m,  1972. 

Chile:  SIO  72-184  (3,  35-39  HL,  182-205+  TL),  18°40.4'S, 
70°36.0'W,  768-967  m,  THOMAS  WASHINGTON,  field  no. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  5 


TABLE  3.  RANGE,  MEAN  (x),  AND  STANDARD  DEVIATION 
(S.D.)  OF  SELECTED  MEASUREMENTS  (VALUES  IN  PERCENT- 
AGE OF  HEAD  LENGTH)  AND  COUNTS  OF  Two  SUBSPECIES  OF 

NEZUMIA  LORICATA  (GARMAN). 


Character 

Sub- 
species 

n 

Range 

x        S.D. 

ID.  rays  (total) 

loricata 

11 

12-14 

12.73      0.8 

atomos 

9 

11-13 

11.78      0.8 

IP.  rays 

loricata 

23 

20-24 

21.48      1.1 

atomos 

18 

21-25 

22.22      1.0 

2P.  rays 

loricata 

24 

10-11 

10.92      0.3 

atomos 

18 

11-12 

11.28      0.5 

GR  I  (total) 

loricata 

12 

9-12 

10.58      0.9 

atomos 

9 

11-14 

12.44      0.9 

GR  II  (total) 

loricata 

12 

10-12 

10.83      0.7 

atomos 

9 

11-13 

12.56      0.7 

Postrostral  len. 

loricata 

10 

70-75 

72.60 

.4 

of  head 

atomos 

9 

67-72 

69.56 

.3 

Snout  length 

loricata 

12 

29-32 

30.75 

.1 

atomos 

9 

32-37 

33.89 

.4 

Internasal 

loricata 

11 

20-24 

22.00 

.2 

length 

atomos 

9 

22-25 

23.56 

.0 

Interorbital 

loricata 

12 

19-24 

21.00 

.4 

width 

atomos 

9 

22-26 

23.78 

.6 

Orbit  to 

loricata 

12 

32-36 

33.92 

.0 

preopercle 

atomos 

9 

34-38 

35.89 

2 

MV72-II-27,  7  May  1972.  —CAS  36807  (6,  27-50  HL,  150+ 
to  287  TL),  24°29.5'S,  70°40'W,  950  m,  ANTON  BRUUN  cr. 
ISA,  sta.  714  (field  no.  LWK.66-60),  16  Aug.  1966.  —CAS 
36810(7,34^*1  HL,  184+ to  220TL),  32°08.5'S,  71°43'W,  960 
m.  ANTON  BRUUN  cr.  ISA,  sta.  703  (field  no.  LWK66-47),  12 
Aug.  1966.  —CAS  36809  (3,  36-^2  HL,  218-220  TL),  32°17'S, 
71°39.5'W,  580  m,  ANTON  BRUUN  cr.  18A,  sta.  702  (field  no. 
LWK66-44),  1 1  Aug.  1966.  —CAS  36805  (28,  29-50  HL,  162- 
300+  TL),  33°39'S,  72°09.5'W,  1,170-1,480  m,  ANTON  BRUUN 
cr.  18A,  sta.  699  (field  no.  LWK66-41),  10  Aug.  1966.  —CAS 
36811  (2,  35-39  HL,  208-224  TL),  34°06.5'S,  72°18.5'W,  750 
m,  ANTON  BRUUN  cr.  ISA,  sta.  687  (field  no.  LWK66-25),  5 
Aug.  1966.  —CAS  36808  (8,  28-39  HL,  148+  to  203  TL), 
34°53.5'S,  72°44'W.  780-925  m,  ANTON  BRUUN  cr.  ISA,  sta. 
698  (field  no.  LWK66-40),  9-10  Aug.  1966. 


Nezumia  loricata  (Garman) 
(Figures  15c,  16,  18g;  Table  3) 

Macrurus  loricatus  GARMAN,  1899:208-209,  pi.  47,  figs.  2-2b 
(original  description;  specimens  from  ALBATROSS  sta.  3409 
and  3410,  off  Galapagos  Islands,  327-331  fms). 

Lionurus  (Nezumia)  loricatus:  GILBERT  AND  HUBBS  1916:146 
(listed). 

Nezumia  loricata:  MAKUSHOK  1967:table  18. 

DIAGNOSIS.— A  species  of  Nezumia  with  10- 
12  (usually  1 1)  pelvic  fin  rays.  Gill-rakers  on  first 
arch  1 1-14  (8-1 1  on  lower  limb);  on  second  arch 
10-13  (8-11  on  lower  limb).  Spinules  on  body 
scales  moderately  to  broadly  lanceolate;  no  en- 
larged middle  row  of  spinules  on  scales.  Barbel 


length  10-17  percent  HL;  upper  jaw  length  29- 
33.  First  dorsal  fin  uniformly  blackish. 

DESCRIPTION. — General  features  of  fish  seen 
in  Figure  15c.  Body  rather  deep,  especially  in 
adults;  greatest  depth  at  origin  of  first  dorsal  fin 
significantly  greater  than  postrostral  length  of 
head.  Head  length  varies  from  17  to  20  percent 
of  total  length.  Rictus  extends  to  below  middle 
of  orbit;  maxillary  extends  to  vertical  slightly 
behind  middle  of  orbit.  Tubercular  scutelike 
scales  at  tip  and  lateral  angles  of  snout  spiny  and 
stout;  the  terminal  scute  bifid.  Interorbital  space 
flat  to  slightly  concave,  relatively  narrow,  1.3- 
1.7  into  orbit.  Gill  membranes  broadly  attached 
across  isthmus  with  little,  if  any,  free  fold.  Gill 
openings  extend  anteroventrally  to  a  point 
slightly  behind  vertical  through  posterior  margin 
of  orbits.  Dermal  window  of  light  organ  small, 
roundish,  situated  between  pelvic  fin  bases. 
Anus  much  closer  to  origin  of  pelvic  fin  than  to 
origin  of  anal  fin.  Gill  filaments  short,  length  of 
longest  less  than  half  diameter  of  orbit. 

Second  spinous  ray  of  dorsal  fin  stout,  armed 
on  leading  edge  with  sharp,  pointed,  reclined 
teeth;  tip  of  ray  slightly  produced.  Paired  fins 
moderate  in  size;  outer  ray  of  pelvic  fin  extends 
beyond  first  three  or  four  anal  rays,  other  pelvic 
rays  barely  (or  do  not)  reach  anal  fin  origin. 

Body  scales  relatively  adherent  (compared 
with  those  of  N.  convergens  and  N.  latirostra- 
ta).  Spinules  on  body  scales  dagger-shaped  to 
narrowly  triangular,  densely  cover  exposed 
fields,  arranged  in  longitudinal  series  that  con- 
verge towards  middle  of  field.  Scales  absent  on 
ventral  surface  of  snout  and  posteriorly  along 
part  of  suborbital  region;  naked  areas  dotted 
with  sensory  papillae,  arranged  in  discrete  rows 
and  clusters  in  most  specimens.  Mandibular 
rami  covered  with  small,  thin,  rather  deciduous 
scales. 

Teeth  small,  in  bands  of  moderate  width  in 
both  jaws.  Outer  series  of  teeth  in  upper  jaw 
slightly  enlarged. 

Color  in  alcohol  swarthy  to  medium  brown. 
Ventral  aspects  of  trunk  and  head  much  darker. 
All  fins  blackish.  Oral  cavity  dusky.  Peritoneal 
lining  blackish  and  densely  punctate  in  Chilean 
specimens  (subspecies  atomos),  but  more  flesh 
colored  with  scattered  large  punctations  in  TE 
VEGA  Galapagos  specimens  (subspecies  lorica- 
ta). 

GEOGRAPHIC  VARIATION. — I  here  recognize 
two  subspecies  of  Nezumia  loricata  based  on 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


175 


differences  in  certain  counts,  measurements, 
and  structural  features  enumerated  and  dis- 
cussed below. 

Key  to  the  Subspecies  of  Nezumia  loricata 

la.  Terminal  snout  scute  composed  of  two 
distinctly  separated  halves.  Gill-rakers  on 
first  arch  9-12  total.  Postrostral  length  of 
head  70-75%  of  head  length  (Jc  =  72.6); 
snout  length  26-32%  (JE  =  30.8).  Head 
length  17.6-20.0%  of  total  length  (x  = 

19.0).  Galapagos  Islands 

N.  loricata  loricata 

Ib.  Each  half  of  terminal  snout  scute  closely 
adpressed,  without  a  deep,  median  gap 
between  them.  Gill-rakers  on  first  arch 
1 1-14  total.  Postrostral  length  of  head  67- 
71%  HL  (x  =  69.6);  snout  length  32-37 
(jc  =  33.9).  Head  length  16.6-18.2%  of  to- 
tal length  (jc  =  17.5).  Off  Chile 

N.  loricata  atomos,  new  subspecies 

The  few  specimens  of  restricted  size  repre- 
senting the  populations  off  Chile  and  the  Gala- 
pagos Islands  limit  comparisons,  but  there  ap- 
pear to  be  sufficient  differences  in  the  study 
material  to  warrant  formal  recognition  of  the  two 
populations.  Frequency  distributions  of  counts 
and  measurements  showing  notable  differences 
between  the  subspecies  are  enumerated  in  Table 
3.  Chief  among  the  other  differences  is  the  shape 
of  the  terminal  snout  scute  (Fig.  16) — in  Gala- 
pagos specimens,  the  scute  is  composed  of  two 
coarsely  spined  halves  medially  split  almost  to 
the  base,  whereas  in  Chilean  specimens,  the 
scute  is  rather  finely  spined  with  the  two  halves 
closely  adjoined  medially,  with  little  gap  be- 
tween. The  tubercular  scales  along  the  leading 
edge  of  the  snout  and  at  the  lateral  angles  are 
also  more  coarsely  spined  in  the  Galapagos 
specimens,  and  the  Chilean  specimens  have  a 
broader  double  row  of  thickened  scales  along 
the  suborbital  region. 

The  two  populations  of  N.  loricata  are  widely 
disjunct,  and  apparently,  no  material  from  in- 
tervening areas  has  been  collected  despite  nu- 
merous trawl  hauls  at  appropriate  depths  along 
the  coasts  of  Ecuador,  Peru,  and  Chile. 

The  subspecific  name  atomos  is  derived  from 
the  Greek  word  meaning  indivisible  or  uncut,  in 
reference  to  the  relatively  undivided  terminal 
snout  scute  of  this  subspecies. 

COUNTS  (of  both  subspecies  combined). —  ID. 


a 


FIGURE  16.  Nezumia  loricata  (Garman).  Dorsal  views  of 
terminal  snout  scute  in  two  subspecies:  (a)  N.  loricata  lori- 
cata; (b)  N.  loricata  atomos. 


11,9-12;  IP.  20-25;  2P.  10-12.  Gill-rakers  on  first 
arch  1-3  +  8-11;  on  second  arch  2  +  8-11. 
Scales  below  first  dorsal  9  to  \1Vi;  below  second 
dorsal  ll/2  to  9;  below  midbase  of  first  dorsal  6l/2 
to  ll/2;  over  distance  equal  to  predorsal  length 
of  head  35-41.  Pyloric  caeca  20-32  (jc  =  25.9,  n 
=  9). 

MEASUREMENTS  (of  both  subspecies  com- 
bined, other  than  those  given  in  Table  3). — Total 
length  175+  to  297+  mm,  head  length  34-52. 
The  following  in  percent  of  head  length:  preoral 
length  25-30;  orbit  diameter  31-36;  postorbital 
length  of  head  36-40;  suborbital  width  14-17; 
upper  jaw  length  29-33;  length  pelvic  fin  44-68; 
length  barbel  10-17;  length  outer  gill  slit  14-17; 
preanal  length  140-157;  base  outer  pelvic  ray  to 
anal  origin  31-44;  anus  to  anal  origin  16-28; 
greatest  body  depth  67-87;  height  first  dorsal  fin 
80-101;  length  pectoral  fin  48-63. 

COMPARISONS  AND  RELATIONSHIPS. — Ne- 
zumia loricata  appears  most  closely  related  to 
N.  latirostrata,  but  the  two  are  readily  distin- 
guished by  differences  given  in  the  key  and  in 
the  description  of  N.  latirostrata.  Nezumia  lor- 
icata may  be  distinguished  from  another  close 
relative,  N.  converge ns,  by  differences  in  gill- 
raker  numbers,  spinules  on  body  scales,  relative 
body  depth,  and  placement  of  anus  (see  com- 
parisons in  description  of  N.  convergens).  Other 
eastern  Pacific  species  of  the  genus  are  differ- 
entiated from  N.  loricata  by  characters  given  in 
the  key. 

DISTRIBUTION. — Galapagos  (subspecies  lori- 
cata) and  central  Chile  (subspecies  atomos). 

SIZE.— To  at  least  52  mm  HL  and  300  mm  TL. 

MATERIAL  EXAMINED. — Galapagos  (subspecies  loricata): 
MCZ  28577  (4  syntypes,  45-55  mm  HL,  215+  to  250  mm  TL), 
and  USNM  57859  (1  syntype,  47  HL,  240+  TL),  00°19'N, 
90°34'W,  331  fms  (605  m),  ALBATROSS  sta.  3410,  3  Apr.  1891. 
—MCZ  28576  (1  syntype,  48  HL,  260  TL),  00°18'40"N, 
90°34'W,  327  fms  (598  m),  ALBATROSS  sta.  3409,  3  Apr,  1881. 


176 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  5 


FIGURE  17.     Nezumia  parini  Hubbs  and  Iwamoto,  1977.  Paratype,  CAS  29414,  35  mm  HL,  216  mm  TL,  from  off  Peru. 


— USNM  135603  (remains  of  about  6  specimens,  mostly  dis- 
integrated), 00°29'S,  89°54'30"W,  392  fms  (717  m),  ALBATROSS 
sta.  2818,  15  Apr.  1888.  —CAS  38329  (7,  34-50  HL,  175+  to 
250  TL),  1°06'S,  89°22'W,  700-800  m,  TE  VEGA  cr.  19,  sta. 
102,  12  Aug.  1968.  — CAS-SU  25246  (2,  44^t6  HL,  poor  con- 
dition), captured  "between  Galapagos  and  Panama,  ALBA- 
TROSS 1888." 

Chile  (subspecies  atomos):  HOLOTYPE,  CAS  38312  (51  mm 
HL,  297+  mm  TL)  and  8  PARATYPES,  CAS  43427  (47-51  HL, 
270-290  TL),  33°39'S,  72°09'W,  1,170-1,480  m,  ANTON 
BRUUN  cr.  18A  (field  no.  LWK66^tl),  10  Aug.  1966. 


Nezumia  parini  Hubbs  and  Iwamoto 

(Figures  17,  18o) 

Nezumia  parini  HUBBS  AND  IWAMOTO,  1977:246-250,  fig.  8, 
11,  table  4. 

DIAGNOSIS. — A  bathypelagic  species  of  Ne- 
zumia with  11-12  pelvic  fin  rays.  First  and  sec- 
ond gill  arches  each  with  10-13  rakers.  Small, 
circular  body  scales  each  with  1-15  long,  erect, 
needlelike  spinules.  Scale  rows  below  second 
dorsal  about  11-14.  Barbel  length  7-13  percent 
HL;  upper  jaw  length  31-36  percent  HL;  outer 
pelvic  fin  ray  length  71-160  percent  HL.  Overall 
color  dark  brown  to  black. 

REMARKS. — General  features  of  the  fish  can 
be  seen  in  Figure  17.  This  distinctive  species, 
described  in  detail  in  the  original  description,  is 
the  only  Nezumia  with  bathypelagic  adults,  al- 
though in  other  species  (e.g.,  N.  stelgidolepis) 
the  early-life  stages  are  probably  bathypelagic. 
The  relationships  of  N.  parini  are  obscure,  but 
probably  lie  closest  to  the  convergens-latiros- 
trata-loricata-orbitalis  complex  of  species.  The 
dense,  fine,  erect  spinules  on  the  small  scales 
give  a  distinctive  furry  texture  and  feel  to  body 
surfaces. 

SIZE. — Nezumia  parini  is  a  small  species  at- 


taining a  head  length  of  35  mm  and  a  total  length 
of  216  mm. 

DISTRIBUTION. — Midwaters  of  the  equatorial 
eastern  Pacific  and  off  the  western  shores  of  the 
Americas  between  Panama  (7°30'N)  and  north- 
ern Chile  (20°00'S). 

MATERIAL  EXAMINED. — See  original  description.  In  addi- 
tion, one  specimen  recently  discovered  in  the  CAS-SU  col- 
lections filed  under  unidentified  Gadidae:  CAS-SU  57651  (for- 
merly New  York  Zoological  Society  cat.  no.  28704),  (1,  35.6 
mm  HL,  202  mm  TL),  off  Colombia,  4°45'N,  78°02'W,  meter 
net  fished  over  bottom  depth  of  500  fms  (914  m),  ZACA  sta. 
233,  3  Apr.  1938. 

ACKNOWLEDGMENTS 

I  thank  the  many  persons  who  helped  in  dif- 
ferent aspects  of  this  study.  The  following 
helped  in  the  loan  of  specimens  and/or  provided 
information  and  assistance  during  museum  vis- 
its: T.  Arai  (Nippon  Luther  Shingaku  Daigaku, 
Tokyo);  N.  Chirichigno-F.  (IMARPE);  D.  M. 
Cohen  (National  Marine  Fisheries  Service, 
Washington,  D.C.);  C.  E.  Dawson  (GCRL);  D. 
Hubendick  (Natural  History  Museum,  Gote- 
borg);  C.  L.  Hubbs  (deceased),  R.  H.  Rosenblatt 
and  assistants  (SIO);  C.  Karrer  (formerly  ISH); 
R.  J.  Lavenberg  and  assistants  (LACM);  R.  Lee 
(University  of  Alaska);  K.  Leim  and  K.  Hartel 
(MCZ);  T.  McLellan  (formerly  MCZ  and  Mas- 
sachusetts Institute  of  Technology);  C.  R.  Rob- 
ins (UMML);  C.  L.  Smith  and  G.  Nelson 
(AMNH);  V.  G.  Springer  and  assistants 
(USNM);  M.  Stehman  (ISH) 

W.  N.  Eschmeyer  examined  and  photo- 
graphed the  holotype  of  Macruroplus  potronus, 
critically  reviewed  the  manuscript,  and  provided 
useful  advice  during  the  study.  L.  J.  Dempster 
assisted  in  literature  searches,  reviewed  the 


IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


177 


FIGURE  18.     Otoliths  (left  saggita)  from  (a)  Nezumia  parini, 
(b)  N.  pulchella,  (c)  N.  liolepis,  (d)  Ventrifossa  nigromacu- 
lata,  (e)  Nezumia  latirostrata,  (/)  N.  convergens,  (g)  N.  lor- 
icataatomos,  (h)N.pudens,  and(i)N.stelgidolepis.  Increments 
on  scale  bars  equal  1  mm. 

manuscript,  and  advised  on  grammatical,  no- 
menclatural,  and  editorial  matters.  O.  Okamura 
reviewed  the  manuscript,  offered  advice  and  in- 
formation, and  provided  many  profitable  hours 
of  discussions  on  the  Macrouridae.  K.  P.  Smith 
rendered  the  fine  illustrations  of  Mataeocepha- 
lus  tenuicauda  (Fig.  8)  and  Nezumia  ventralis 
(Fig.  14b).  Curatorial  and  technical  assistance 
was  provided  by  J.  E.  Gordon,  T.  Keating.  B. 


Powell,  W.  C.  Ruark,  and  P.  Sonoda.  My  sin- 
cerest  thanks  to  all. 

This  study  was  supported  in  part  by  a  grant 
from  the  National  Science  Foundation  (BMS 
075-03153),  Tomio  Iwamoto,  principal  investi- 
gator. 

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IWAMOTO:  EASTERN  PACIFIC  MACROURIDAE 


179 


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pp. 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  6,  pp.  181-227,  17  figs.  July  2,  1980 


REVISION  OF  THE  EASTERN  PACIFIC  SYNGNATHIDAE 

(PISCES:SYNGNATHIFORMES),  INCLUDING 

BOTH  RECENT  AND  FOSSIL  FORMS 

By 
Ronald  A.  Fritzsche 

Department  of  Biology,  The  University  of  Mississippi, 
University,  Mississippi  38677 


Abstract:  The  marine  and  estuarine  eastern  Pacific  Syngnathidae  comprises  17  extant  and  3  extinct  species. 
Diagnostic  characters  for  species  and  genera,  including  meristic  and  morphometric  characters,  and  osteological 
features,  are  given.  All  species  both  living  and  fossil  are  diagnosed  and  described. 

The  recognized  species  and  their  ranges  are:  Hippocampus  ingens  Girard  [=//.  hildebrandi]  (San  Francisco 
Bay,  California,  south  to  Pucusana,  Peru,  including  the  Gulf  of  California);  Doryrhamphus  melanopleura  (Bleek- 
er)  (Indo-Pacific;  in  the  eastern  Pacific  from  Bahia  Magdalena,  Baja  California,  south  to  Isla  la  Plata,  Ecuador, 
including  the  Gulf  of  California,  Galapagos  Islands,  and  Clipperton  Island);  Doryrhamphus  paulus  n.sp.  (Islas 
Revillagigedo,  Mexico);  Leptonotus  blainvilleanus  (Eydoux  and  Gervais)  [=5.  acicularis]  (Hornitos,  Chile,  to 
Golfo  Nuevo,  Argentina);  Bryx  arctus  (Jenkins  and  Evermann)  (Tomales  Bay,  California,  south  to  Mazatlan, 
Mexico,  including  the  Gulf  of  California);  Bryx  heraldi  n.sp.  (Islas  Juan  Fernandez  and  Isla  San  Felix,  Chile); 
Bryx  coccineus  (Herald)  (Bahia  Banderas,  Mexico,  south  to  Punta  Aguja,  Peru,  and  the  Galapagos  Islands); 
Bryx  veleronis  Herald  (Galapagos  Islands;  Islas  Revillagigedo;  Isla  Murcielago  and  Isla  del  Cario,  Costa  Rica; 
and  Islas  San  Jose  and  Canal  de  Afuera,  Panama);  lir\\  clarionensis  n.sp.  (Isla  Clarion,  Mexico);  Syngnathus 
auliscus  (Swain)  (Santa  Barbara  Channel,  California,  south  to  Paita,  Peru,  including  the  Gulf  of  California); 
Syngnathus  carinatus  (Gilbert)  (confined  to  upper  Gulf  of  California);  Syngnathus  exilis  (Osburn  and  Nichols) 
(Half  Moon  Bay,  California,  to  Bahia  Magdalena,  Baja  California,  and  Isla  Guadalupe,  Mexico);  Syngnathus 
californiensis  Storer  (Bodega  Bay,  California,  south  to  Bahia  Santa  Maria,  Baja  California);  Syngnathus  ma- 
crobrachium  n.sp.  (Tumbes,  Peru,  south  to  Puerto  Montt,  Chile);  Syngnathus  euchrous  n.sp.  (Redondo  Beach, 
California,  to  Punta  Eugenia,  Baja  California);  Syngnathus  leptorhynchus  Girard  [=S.  griseolineatus ,  S.  bar- 
barae]  (southeastern  Alaska  south  to  Bahia  Santa  Maria,  Baja  California);  Syngnathus  insulae  n.sp.  (Isla 
Guadalupe,  Mexico). 

Studies  of  growth  and  variation  show  that  S.  leptorhynchus  is  highly  variable  with  each  population  distinct. 
Marked  seasonal  variation,  when  combined  with  growth  data,  indicates  that  individuals  probably  live  for  one 
year  or  less. 

The  fossil  pipefishes  of  Southern  California  are  all  only  known  from  the  Miocene.  Hipposyngnathus  imporcitor 
n.sp.  from  the  upper  Modelo  Formation  is  most  closely  related  to  two  species  from  the  Oligocene  of  Europe. 
Syngnathus  emeritus  n.sp.  is  known  only  from  the  Puente  Formation.  Syngnathus  avus  Jordan  and  Gilbert  is 
known  from  the  lower  Modelo  Formation. 

A  comparison  between  inferred  relationships  of  the  various  species  and  their  geographical  distribution  sug- 
gests that  the  evolution  of  the  eastern  Pacific  Syngnathus  is  a  result  of  at  least  two  separate  invasions.  Doryr- 
hamphus melanopleura  invaded  the  eastern  Pacific  by  crossing  the  East  Pacific  Barrier  and  gave  rise  to  Doryr- 
hamphus paulus.  Leptonotus  blainvilleanus  is  related  to  other  species  of  Leptonotus  in  New  Zealand  and  southern 
Australia  and  was  probably  derived  from  an  ancestor  in  these  areas  via  waif  dispersal. 

The  reduction  and  loss  of  elements  of  the  branchial  skeleton  is  useful  in  characterizing  urophorine  genera 
and  may  be  of  general  use  when  relationships  within  the  family  Syngnathidae  are  studied  in  more  detail. 

181 


182 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


INTRODUCTION 

Pipefishes  of  the  family  Syngnathidae  inhabit 
most  temperate  and  tropical  seas.  In  the  eastern 
Pacific,  pipefishes  occur  from  southeastern 
Alaska  to  Tierra  del  Fuego.  They  are  primarily 
marine  or  euryhaline,  but  some  species  are  con- 
fined to  fresh  water. 

Even  though  the  family  Syngnathidae  con- 
tains the  seahorses,  of  general  interest  to  aquar- 
ists,  the  family  has,  in  general,  been  poorly  stud- 
ied. There  are  a  number  of  undescribed  species 
and  the  intrafamilial  relationships  are  poorly 
understood,  due,  in  part,  to  the  great  variability 
in  meristic  and  morphometric  characters. 

A  revision  of  the  Syngnathidae  was  published 
by  Duncker  (1915),  but  the  American  species 
were  not  included.  Ginsburg  (1937)  and  Herald 
(1940-1965)  have  been  the  major  contributors  to 
the  taxonomy  of  the  American  syngnathids.  To 
date,  a  definitive  treatment  of  the  eastern  Pacific 
syngnathids  has  not  appeared. 

Detailed  osteological  work  on  adult  syngnath- 
ids is  limited  to  that  of  Jungersen  (1910),  Rauther 
(1925),  and  Banister  (1967).  Jungersen's  study 
included  the  genera  Hippocampus,  Syngnathus, 
and  Nerophis,  two  of  which  occur  in  the  eastern 
Pacific.  Rauther  also  studied  the  osteology  of 
Syngnathus,  as  did  Banister.  The  osteology  of 
the  other  four  genera  of  eastern  Pacific  syng- 
nathids has  not  been  treated.  It  is  generally  rec- 
ognized that  for  the  study  of  higher  taxa,  osteo- 
logical characters  are  a  good  indicator  of 
phylogeny  because  of  their  conservative  nature. 

The  goal  of  this  study  has  been  to  characterize 
the  eastern  Pacific  genera  and  species  of  Syng- 
nathidae, both  fossil  and  recent,  and  to  examine 
intraspecific  variation.  In  addition,  a  preliminary 
examination  of  the  osteology  of  a  few  of  the 
syngnathid  genera  was  made  to  help  in  under- 
standing the  relationships  between  at  least  a  few 
of  the  genera,  and  as  a  test  of  the  relationships 
proposed  by  Herald  (1959)  based  on  the  method 
of  brood-pouch  closure. 

MATERIALS  AND  METHODS 

Specimens  that  have  contributed  to  the  data 
are  listed  in  the  Material  Examined  section  for 
each  species.  In  those  lists,  the  number  of  spec- 
imens is  given,  followed  by  the  size  range  (SL 
in  mm)  enclosed  in  parentheses.  If  the  size  range 
was  not  determined  or  the  specimens  are  dam- 
aged, the  range  of  standard  length  is  not  given. 


Abbreviations  for  listed  collections  are:  AMS, 
the  Australian  Museum,  Sydney;  BC,  Univer- 
sity of  British  Columbia:  BMNH,  British  Mu- 
seum (Natural  History);  BOC,  Bingham  Ocean- 
ographic  Collection,  Yale  University;  CAS, 
California  Academy  of  Sciences;  EMBCh,  Es- 
tacion  de  Biologfa  Marina,  Chile;  GCRL,  Gulf 
Coast  Research  Laboratory  Museum;  HSU, 
Humboldt  State  University;  IMARPE,  Institute 
del  Mar,  Peru;  LACM,  Natural  History  Museum 
of  Los  Angeles  County;  MCZ,  Museum  of  Com- 
parative Zoology,  Harvard  University;  MNHN, 
Museum  National  d'Histoire  Naturelle,  Paris; 
MNMH  P,  Museo  Nacional  de  Historia  Natural, 
Santiago,  Chile;  SCCWRP,  Southern  California 
Coastal  Water  Research  Project;  SIO,  Scripps 
Institution  of  Oceanography  Marine  Vertebrates 
Collection;  SU,  Stanford  University  (now 
housed  at  CAS);  UCLA,  Department  of  Zoolo- 
gy, University  of  California,  Los  Angeles; 
UMMZ,  University  of  Michigan  Museum  of  Zo- 
ology, USNM,  National  Museum  of  Natural 
History,  Smithsonian  Institution;  UW,  Univer- 
sity of  Washington. 

Measurements  were  made  with  dial  calipers 
to  the  nearest  0.1  mm  for  lengths  less  than  17 
cm;  an  ocular  micrometer  was  used  for  mea- 
surements less  than  2  mm.  Measurements  great- 
er than  17  cm  were  made  to  the  nearest  mm  with 
a  centimeter  rule. 

The  principle  characters  used  in  identifying 
syngnathids  are  illustrated  in  Figures  1  and  2 
(based  on  a  generalized  syngnathid).  The  ter- 
minology used  for  the  various  ridges  on  the  body 
is  that  of  Herald  (1943). 

The  condition  of  the  lateral  trunk  ridge  (Fig. 
2)  is  important  in  characterizing  syngnathid  gen- 
era. It  is  subcontinuous  with  the  superior  tail 
ridge  in  Bryx  and  Syngnathus,  but  continuous 
in  Leptonotus.  In  Hippocampus  and  Doryrham- 
phus  the  lateral  trunk  ridge  is  continuous  with 
the  inferior  tail  ridge.  Clausen  (1956)  has  shown 
that  these  ridge  patterns  are  not  wholly  consis- 
tent within  species,  however,  they  are  of  value 
when  used  with  other  characters. 

Scutella  (Fig.  1),  small  oval  plates  interpolated 
between  the  larger  dermal  plates,  may  be  pres- 
ent or  absent.  Their  size  is  important  in  species 
determinations  and  is  therefore  included  in  the 
species  descriptions.  Size  is  given  as  a  compar- 
ison of  an  individual  scutellum  with  an  adjacent 
plate. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 

r- 


183 


I 

FIGURE  1.  Characters  used  in  syngnathid  identification 
(modified  after  Herald  1943).  (A)  Characters  in  the  region  of 
the  head:  a.  snout  ridge;  b.  supraorbital  ridge;  c.  supraocci- 
pital,  nuchal,  and  prenuchal  ridges;  d.  opercular  ridge;  e.  pec- 
toral cover  plate  ridges;/,  ventral  trunk  ridge.  (B)  Characters 
in  the  region  of  the  dorsal  fin:  g.  superior  trunk  ridge;  h. 
superior  tail  ridge;  i.  lateral  trunk  ridge;  j.  inferior  trunk  ridge; 
k.  anus;  /.  anal  fin;  m.  inferior  tail  ridge;  n.  scutellum. 


A  ring  is  defined  as  one  unit  in  the  series  of 
dermal  plates  which  form  definite  bands  around 
the  body. 

Pouch  protecting  plates  are  not  illustrated. 
They  are  merely  ventral  extensions  of  the  infe- 
rior trunk  or  tail  ridge  which  support  the  brood 
pouch  and  protect  the  eggs. 

The  pectoral  cover  plate  is  defined  as  the  plate 
covering  the  base  of  the  pectoral  fin. 

The  methods  of  making  counts  and  measure- 
ments are  those  of  Hubbs  and  Lagler  (1958), 
with  the  following  exceptions: 

Number  of  trunk  and  tail  rings:  The  ring  bearing 
the  pectoral  fins  is  counted  as  the  first  trunk 
ring;  the  ring  bearing  the  anus  is  the  last  trunk 
ring.  The  ring  bearing  the  anal  fin  is  the  first 
tail  ring.  If  the  anus  and  the  anal  fin  are  borne 
on  the  same  ring,  then  that  ring  is  the  first  tail 
ring  and  the  preceeding  ring  is  the  last  trunk 
ring. 

Number  of  dorsal  and  anal  fin  rays:  last  two  rays 
are  counted  as  two. 

Number  of  rings  covered  by  dorsal  fin:  fractions 


FIGURE  2.  Configuration  of  ridges  above  the  anus  (modi- 
fied after  Herald  1943).  (A)  Lateral  trunk  ridge  subcontinuous 
with  superior  tail  ridge.  (B)  Lateral  trunk  ridge  continuous 
with  superior  tail  ridge.  (C)  Lateral  trunk  ridge  continuous 
with  inferior  tail  ridge. 

of  a  ring  estimated  to  the  nearest  tenth  of  a 
ring. 

Number  of  lateral-line  papillae:  range  of  the 
number  of  lateral-line  papillae  per  dermal 
plate. 

For  Hippocampus  the  standard  length  is  the 
distance  from  the  tip  of  the  coronet,  with  the 
head  held  perpendicular  to  the  trunk,  to  the  tip 
of  the  tail,  with  the  tail  held  straight  (Ginsburg 
1937). 

In  species  diagnoses  the  modal  range  of  a  par- 
ticular meristic  character  is  given  in  parentheses 
after  the  range  of  that  character. 

Osteology  was  studied  from  trypsin-digested 
and  alizarin-stained  specimens  (Taylor  1967) 
listed  below.  Osteological  nomenclature  follows 
Jungersen  (1910)  and  Banister  (1967).  Bones 
were  drawn  with  the  aid  of  a  camera  lucida. 

Diagnoses  are  based  on  adults  unless  other- 
wise noted. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


Statistical  procedures  follow  the  text  of  Dixon 
and  Massey  (1957)  except  that  the  regression 
analyses  are  based  on  the  Bartlett  regression 
analysis  as  given  by  Hoese  (1971).  Unless  oth- 
erwise specified,  differences  are  considered  sig- 
nificant ai  P  ^  .05. 


basisphenoid;  no  metapterygoid;  eggs  incubated 
by  males  in  a  special  area  under  the  trunk  or 
tail,  which  may  or  may  not  be  developed  into  a 
pouch;  no  pyloric  sphincter;  no  distinct  stom- 
ach; right  kidney  only  present,  aglomerular;  pre- 
dorsals  reduced  to  2-3  nuchal  plates. 

The  family  is  usually  divided  into  two  groups: 
Gastrophori — those  which  develop  the  brood 
pouch  under  the  abdomen;  Urophori — those 
with  the  brood  pouch  under  the  tail. 


CLEARED  AND  STAINED  MATERIAL  EXAMINED. — Heraldia 
nocturna,  SIO  75-54,  1,  Sydney,  Australia;  Maroubra  perser- 
rata,  SIO  75-53,  1,  Sydney,  Australia;  Choeroichthys  brachy- 
soma  SIO  73-196,  1,  Timor  Sea;  Doryrhamphus  melanopleu- 
ra,  SIO  65-343,  1,  Gulf  of  California;  Dunckerocampus 

dactylophorus,  CAS  uncat.,  1,  unknown;  Dent/rostrum  jans-  Key  tO  Genera  of  Eastern  Pacific 

si,  CAS  14148,  3  paratypes,  Thailand;  Syngnathoides  biacu-  Svnenathidae 

leatus,  SIO  61-693,  1,  10°N,  103°50'E;  Leptonotus  blainvil- 

leanus,  USNM  176569,  1,  isia  Chiloe,  Chile;  Leptonotus  la.  Caudal  fin  absent;  tail  prehensile;  head  at 

blainvilleanus,  USNM  176564,  1,  Bahia  Lin,  Chile;  Parasyng-  right  angle  to  main  body  axis 

nathus  elucens,  CAS  13696,  1,  Virgin  Islands;  Ichthyocampus  HlDDOCamDUS  RafinCSQUC 

belcheri,  CAS  uncat.,  2,  Philippine  Is.;  Micrognathus  brevi-  ../I  i  /.  .    •,  •• 

rostris,  SIO  73-196,  4,  Timor  Sea;  Penetopteryx  taenio-  lb'  Caudal  fi.n  PreS6nt'  tal1  neV6r  Pensile; 
cephalus,  SIO  66-587,  1,  Great  Tulear  Reef;  Syngnathus  can-  head  in  line  with  main  body  axis  __ 

natus,  S.  Guevarra  pers  coll.,  1,  Gulf  of  California;  S.  acus,  2SL.    Brood  pouch  Under  abdomen     3 

SIO  73-310,  1,  Yugoslavia;  S.  auliscus,  SIO  68-168,  1,  Sonora,  2b     Brood  pouch  Under  tail  4 

Mexico;  S  ca/yor*««5   SIO  H47-180J    Santa  Cmz  Island:  3a' Trunk    rj  more    numerous"" than  "tail 

S.  leptorhynchus,  E.  B.  Brothers  pers.  coll.,  1,  Mission  Bay; 
5.  pelagicus,  SIO  65-358,  1,  western  Atlantic;  Bryx  veleronis,  nngS___  ..  Doryrhamphus  Kaup 

SIO  71-52,  1,  Panama;  B.  clarionensis,  SIO  74-116,  1,  para-  3b.   Trunk  rings  fewer  than  tail  rings    

type,  Clarion  Is.;  B.  dunckeri,  SIO  70-376,   1,  Panama;  B.  OostethliS  Hubbs 

arctus,  SIO  H52-218,  1,  Baja  California;  B.coccineus,  USNM  4a  Lateral  trunk  ridge  Continuous  with  SU- 
220972,  1,  Galapagos  Is.;  Cosmocampus  brachy cephalus,  .  ..  ..  ,  ,  ,  ... 

CAS  24025,  2,  Panama;   Corythoichthys  flavofasciatus ,   R.  P6riOr    tai1    "^    brood    pouch    Without 

Nolan  pers.  coll.,  1,  Eniwetak;  Cory thoichy thy s  sp.,  SIO  73-  protecting  plates;  mature  females  with 

206,  1,  Timor  Sea,  Pseudophallus  starksi,  USNM  208371,  2,  deep,  Compressed  trunk  

Panama;  P.  elcapitanensis,  USNM  208369,  1,  Panama;  Hip-  Leptonotus  Kaup 

pocampus  knda,  SIO-60-250  1  Hawaii,  H.  ingen,  NMFS  4b  LaterarVrunk"ridge""subcontinUOUS  with 
uncat.,  1,  eastern  Pacific;  Pnyllopteryx  jonatus,  SIO  73-361,  .  . 

1,  Australia  superior  tail  ridge;  brood  pouch  with  pro- 

tecting plates;  mature  females  with  sub- 
SYSTEMATICS  cylindrical  trunk  5 

Family  Syngnathidae  Bonaparte,  1838  5a"  DorS^ra,ySA3  °r  fewer'  Sn?Ut  sl!°rt'  C°n' 

tamed  2.3-4.0  times  in  head;  anal  fin  pres- 

Type-Genus:  Syngnathus  Linnaeus,  1758.  ent   Qr  absent;    smaH5    never  longer  than 

DIAGNOSIS. — Syngnathiforms  with  body  en-  124  mm  SL Bryx  Herald 

cased  in  armor  formed  of  dermal  plates  arranged  5b.  Dorsal  rays  26  or  more;  snout  longer,  con- 
in  rings;  pelvic  and  spinous  dorsal  fins  absent;  tained  1.6-2.8  times  in  head;  anal  fin  pres- 
dorsal  and  pectoral  fins  moderately  developed  ent;  larger,  most  species  reach  200  mm  or 

to  absent;  caudal  and  anal  fins  weakly  developed  more  SL Syngnathus  Linnaeus 

to  absent  (tail  often  prehensile  when  caudal  fin 

absent);  opercular  slit  reduced  to  a  small  open-  Records  of  Doubtful  Validity 

ing  at  dorsoposterior  margin  of  opercle,  four         Regan  (1908)  included  Oostethus  brachyurus 

complete  gill  arches  bearing  lobate  gills;  pseu-  (Bleeker)  and  Syngnathus  spicifer  Riippell  in  his 

dobranchiae  present;  supracleithra  and  post-  list  of  the  fishes  occurring  at  Tehuantepec,  Mex- 

cleithra  absent;  ribs  absent;  teeth  lacking  on  ico.  Herald  (1940)  repeated  these  records  but 

jaws,  but  premaxillae  and  dentaries  may  bear  doubted  their  validity.  Herald  (1943)  indicated 

odontoid  processes  (Dawson  and  Fritzsche  that  there  was  considerable  doubt  that  these  two 

1975);  pharyngobranchial  tooth  plates  present  or  species  were  collected  at  Tehuantepec,  because 

absent;  upper  part  of  cleithrum  forms  part  of  he  had  been  able  to  confirm  that  Regan's  spec- 

the  external  armor;  posttemporals  suturally  imens  had  come  from  an  animal  dealer.  Duncker 

united  to  neurocranium;  1-3  branchiostegals;  no  (1915)  had  also  realized  that  the  specimens  had 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


185 


come  from  an  animal  dealer,  but  the  records 
have  persisted.  The  animal  dealer  had  received 
specimens  from  both  Mexico  and  the  Philip- 
pines. O.  brachyurus  and  S.  spicifer  occur  in 
the  Philippines,  and  it  seems  most  probable  that 
the  specimens  were  collected  there. 

There  is,  however,  a  valid  record  of  Oostethus 
from  the  eastern  Pacific.  McCosker  and  Dawson 
(1975)  reported  a  single  individual  of  the  eury- 
haline  Atlantic  Oostethus  lineatus  collected  on 
the  Pacific  side  of  Panama  in  1971  and  concluded 
that  the  specimen  represents  a  transit  of  the  Pan- 
ama Canal. 

Dermatostethus  punctipinnis  Gill  (1863)  was 
described  from  four  specimens  collected  at  San 
Diego.  Presently  there  are  three  type-specimens 
at  the  USNM  (lectotype,  here  designated  as 
USNM  8128;  paralectotypes  here  designated  as 
USNM  214484).  Much  confusion  has  been  gen- 
erated by  these  specimens.  The  types  are  quite 
large  (302-344  mm  SL),  have  a  very  flexible 
"neck,"  and  dark  spotting  on  the  dorsal  fin. 
Herald  (1940,  1941)  referred  D.  punctipinnis  to 
the  synonymy  of  Syngnathus  californiensis  be- 
cause the  latter  is  a  large  species  with  counts 
like  that  of  the  type  material  of  D.  punctipinnis. 
Miller  and  Lea  (1972:212)  suggested  that  D. 
punctipinnis  might  be  a  valid  species,  but  did 
not  include  it  in  their  key  to  California  species. 
I  have  examined  the  types  of  D.  punctipinnis 
and  have  compared  them  to  specimens  of  S. 
acus  from  Europe  and  have  been  unable  to  find 
significant  differences.  I  therefore  relegate  D. 
punctipinnis  to  the  synonymy  of  S.  acus. 

Syngnathus  acus  has  been  recorded  from  the 
Indo- Pacific  (Weber  and  De  Beaufort  1922),  and 
my  examination  of  a  specimen  from  that  region 
(Pakoi,  China)  suggests  the  existence  of  a  dis- 
tinct acus-like  species.  But  the  putative  Califor- 
nia material  is  typical  of  acus  and  could  not  rep- 
resent trans-Pacific  migration. 

Dumeril  (1870)  described  Syngnathus  bairdi- 
anus  from  a  locality  given  as  "Cote  du  Mexique, 
voisine  de  la  Californie."  The  locality  as  listed 
in  the  catalog  of  the  MNHN  is  "Mexique  pres 
la  Californie,  Lucas  1867."  I  examined  the  two 
types  (lectotype,  here  designated  as  MNHN 
6112;  paralectotype,  here  designated  as  MNHN 
2756)  and  found  them  to  be  indistinguishable 
from  S.  pelagicus  Linnaeus.  I  therefore  relegate 
S.  bairdianus  to  the  synonymy  of  S.  pelagicus. 
S.  pelagicus  inhabits  the  Sargasso  Sea  and  ap- 
parently can  be  transported  over  long  distances. 


It  has  been  recorded  from  Tierra  del  Fuego 
(Fowler  1944)  and  from  New  Zealand  (Weber 
and  De  Beaufort  1922).  His  trio  histro,  another 
inhabitant  of  the  Sargasso  Sea,  is  widely  distrib- 
uted and  has  been  recorded  from  the  Galapagos 
Islands  (Schultz  1957).  It  is  therefore  remotely 
possible  that  S.  pelagicus  could  have  been  col- 
lected near  the  coast  of  Mexico. 

None  of  the  above  species  will  be  treated  fur- 
ther because  of  their  doubtful  standing  as  mem- 
bers of  the  eastern  Pacific  fauna. 

Hippocampus  Rafinesque 

Hippocampus  RAFINESQUE,  1810:18  (type-species  by  mono- 
typy,  H.  heptagonus  Rafinesque  [=Syngnathus  hippocam- 
pus Linnaeus]);  DUNCKER  1912:237  (diagnosis);  1915:115 
(diagnosis);  GINSBURG  1937:525  (diagnosis;  discussion). 

Farlapiscus  WHITLEY,  1931:313  (type-species  by  original  des- 
ignation, Hippocampus  breviceps  Peters). 

Hippohystrix  WHITLEY,  1940:44  (type-species  by  original  des- 
ignation, Hippocampus  spinosissimus  Weber). 

DIAGNOSIS. — Urophori  characterized  by  a 
prehensile  tail;  absence  of  caudal  fin,  scutella, 
brood  pouch  protecting  plates,  and  basibranchi- 
als;  head  at  right  angle  to  trunk;  brood  pouch 
sealed  along  midline  except  for  small  anterior 
opening;  occiput  raised  to  form  coronet;  dorsal 
fin  base  raised. 

DISTRIBUTION. — Marine;  world- wide  in  trop- 
ical and  subtropical  seas. 

Hippocampus  ingens  Girard 

(Figures  3  &  7C) 

Hippocampus  ingens  GIRARD,  1859:342  (original  description; 
San  Diego,  California)  (lectotype  here  designated  as  USNM 
982);  JORDAN  AND  GILBERT  1880:23  (San  Diego);  1881:453 
(San  Diego);  JORDAN  AND  JOUY  1881:1  (California);  JOR- 
DAN AND  GILBERT  1882:69  (San  Diego);  1883:386  (descrip- 
tion; range);  EVERMANN  AND  JENKINS  1891:127,  135(Guay- 
mas,  Mexico);  GILBERT  1891:450  (ALBATROSS  sta.  2795); 
EIGENMANN  AND  EIGENMANN  1892: 144  (San  Diego);  VAIL- 
LANT  1894:  70  (Gulf  of  California);  JORDAN  1895:417  (de- 
scription; Mazatlan,  Mexico);  JORDAN  AND  EVERMANN 
1896:776  (synonymy;  description);  GILBERT  AND  STARKS 
1904:57  (Panama  Bay);  GILL  1905:807  (range);  STARKS  AND 
MORRIS  1907:186  (range);  OSBURN  AND  NICHOLS  1916:155 
(Concepcion  Bay);  NICHOLS  AND  MURPHY  1922:506  (Peru); 
MEEK  AND  HILDEBRAND  1923:256  (description;  synony- 
my); BREDER  1928:23  (Cape  San  Lucas,  Concepcion  Bay); 
ULREY  AND  GREELEY  1928:41  (synonymy;  range);  ULREY 
1929:6  (Lower  California,  Gulf  of  California);  1932:77  (Baja 
California);  GINSBURG  1937:534  (range;  synonymy;  descrip- 
tion); ATZ  1937:62  (size);  FOWLER  1944:496  (range); 
KOEPCKE  1962:200  (references;  range);  CLEMENS  AND 
NOWELL  1963:262  (off  Mexico;  in  stomachs  offish);  HUBBS 
AND  HINTON  1963:12  (California  record;  range);  CHIRI- 
CHiGNO-F.  1963:8,  34  (Peru;  range);  CASTRO-AGUIRRE  ET 
AL.  1970:132  (common  in  Gulf  of  California);  MILLER  AND 


186 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


FIGURE  3.  Hippocampus  ingens,  MCZ  35292,  collected  in 
1860  at  San  Francisco,  California,  by  Alexander  Agassiz  (not 
examined). 


LEA  1972:89  (description;  range);  CHIRICHIGNO-F.  1974:  86, 
337  (Peru;  in  key;  range). 

Hippocampus  gracilis  GILL,  1863:282  (original  description; 
Cape  San  Lucas,  Baja  California). 

Hippocampus  ecuadorensis  FOWLER,  1921:446  (original  de- 
scription; Bahfa,  Ecuador). 

Hippocampus  hildebrandi  GINSBURG,  1933:562  (original  de- 
scription; Chame  Point,  Panama);  holotype  USNM  82063); 
1937:579  (description;  range). 

DIAGNOSIS. — A  Hippocampus  with  17-22(19) 
dorsal  rays;  15-17  pectoral  rays;  4-5  anal  rays; 
11-12  (11)  trunk  rings;  37-41  (39)  tail  rings;  dor- 
sal fin  on  1.2-2  +  1.5-4  rings;  total  rings  cov- 
ered by  dorsal  fin  3-4;  brood  pouch  on  5-8  rings; 
head  4.2-5.8  in  SL;  dorsal  fin  base  1.8-2.9  in 


head;  snout  1.8-2.5  in  head;  maximum  size  ex- 
amined 247  mm  SL;  smallest  mature  male  54 
mm,  said  to  attain  12"  (297  mm)  (Miller  and  Lea 
1972),  which  is  approximately  the  size  of  my 
largest  specimen  if  the  head  is  included  in  the 
measurement. 

DESCRIPTION. — Body  ridges  typically  well  de- 
veloped with  a  blunt  recurved  tubercle  at  center 
of  each  plate.  First,  fourth,  seventh,  and  tenth 
trunk  rings  usually  with  larger  and  better  devel- 
oped tubercles.  Sixth,  tenth,  fourteenth,  and 
eighteenth  tail  rings  may  have  strongly  devel- 
oped tubercles,  but  are  usually  more  obsoles- 
cent than  trunk  tubercles.  Coronet  with  five  ra- 
dially arranged  tubercles.  Males  with  the  more 
weakly  developed  tubercles.  Tubercles  in  both 
sexes  generally  become  obsolete  with  growth. 
Strong  nuchal  ridge,  without  spines.  Opercle 
with  faint  radiating  striae.  No  prominent  snout 
ridge.  Internasal  spine  blunt.  Prominent  broad- 
based  supraopercular  spine  directed  laterally. 
Lateral  trunk  ridge  and  superior  tail  ridges  over- 
lap for  one  ring.  Trunk  rings  octangular  under 
dorsal  fin,  heptangular  anteriorly.  Trunk  mod- 
erately compressed.  First  tail  ring  heptangular, 
remainder  quadrangular.  Superior  trunk  and  tail 
ridges  overlap  for  two  to  three  rings.  Dermal 
flaps,  when  present,  on  enlarged  tubercles  and 
head  ridges,  consisting  of  a  stout  base  with  nu- 
merous filamentous  outgrowths. 

Coloration  in  life.  Red,  yellow,  or  green.  One 
specimen  captured  at  La  Paz,  Baja  California, 
had  undersides  and  tips  of  tubercles  yellowish, 
most  of  body  mottled  with  dark  brown  to  black, 
and  covered  with  many  small  dark  spots  as  well 
as  smaller  white  ones.  The  white  spots  tend  to 
coalesce  into  longitudinal  streaks;  yellowish  col- 
oration more  pronounced  on  underside  of  tail. 
White  bands  around  body  every  six  or  seven 
rings.  Dorsal  fin  with  distinct  dark  submarginal 
band.  Pectoral  fins  hyaline. 

Coloration  in  alcohol.  Enlarged  tubercles  usu- 
ally whitish,  often  with  white  ring  around  body 
at  that  point.  Ground  color  dark  brown  with 
small  white  papillae  often  forming  streaks  and 
reticulations,  and  radiating  lines  around  orbit. 
Ventral  surface  of  tail  without  white  markings. 
Dorsal  fin  with  a  dark  band  near  margin.  Median 
abdominal  ridge  often  dark  brown  in  males. 
However,  color  variable;  some  specimens  may 
be  uniform  light  tan. 

HABITAT. — Collections  of  H.  ingens  are  un- 
common. Some  have  been  made  in  shallow 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


187 


water,  but  most  specimens  have  been  captured 
at  depth  with  dredges  or  trawls,  or  at  the  surface 
in  the  open  ocean.  Dredge  and  trawl  collections 
have  usually  been  made  at  depths  of  10  m  or 
more.  Juveniles  (ca.  40  mm)  and  larger  individ- 
uals (115  mm)  are  not  uncommonly  taken  at  the 
surface.  Alverson  (1963)  studied  the  food  items 
in  stomachs  of  eastern  Pacific  yellowfin  tuna, 
Thunnus  albacares,  and  found  H.  ingens  in  18 
stomachs  out  of  the  2846  he  examined.  Judging 
from  the  small  displacement  volumes  of  these 
fish,  they  were  all  juveniles.  One  34-mm  indi- 
vidual (SIO  71-186)  was  taken  from  the  gut  of  a 
bluefin  tuna,  Thunnus  thynnus.  The  habitat  is 
not  yet  precisely  known.  H.  ingens  appears  to 
spend  much  of  its  life  in  the  open  sea. 

COMPARISONS. — Ginsburg  (1933)  described 
H.  hildebrandi  from  Chame  Point,  Panama,  as 
differing  from  H.  ingens  in  having  lower,  broad- 
er tubercles.  My  examination  of  38  specimens 
from  the  eastern  Pacific  has  shown  that  all  spec- 
imens are  referrable  to  H.  ingens.  I  have  ex- 
amined the  types  of  H.  ingens  and  H.  hilde- 
brandi and  conclude  that  the  types  of  H. 
hildebrandi  are  juveniles  of  H.  ingens. 

Since  H.  ingens  is  the  only  species  of  sea- 
horse in  the  eastern  Pacific,  it  is  easily  identifi- 
able. The  closest  relative  of  H.  ingens  is  pos- 
sibly H.  reidi  Ginsburg  from  the  western 
Atlantic,  from  which  it  differs  in  number  of  dor- 
sal rays  (17-22  vs  15-19)  and  in  number  of  tail 
rings  (36-41  vs  34-37).  The  relationship  of  H. 
ingens  to  seahorses  of  the  Indo-Pacific  is  im- 
possible to  determine  because  knowledge  of  the 
genus  Hippocampus  is  very  incomplete. 

RANGE. — San  Francisco  Bay,  California, 
south  to  Pucusana,  Peru,  including  the  Gulf  of 
California.  Infrequently  taken  north  of  central 
Baja  California.  During  periods  of  unusually 
warm  water,  H.  ingens  may  enter  California 
waters. 

MATERIAL  EXAMINED. — California:  San  Diego,  USNM 
982,  1(167),  lectotype:  USNM  214485,  2(125-146),  paralecto- 
types;  Point  Loma,  SIO  63-1085,  1(195). 

Baja  California:  Laguna  Guerrero  Negro,  SIO  61-10,  3(155- 
185).  Bahfa  Sebastian  Vizcaino,  SIO  63-1046,  1(29).  Bahfa 
Magdalena,  SIO  62-713,  1(108);  SIO  64-73,  1(27);  SIO  60-305, 
1(55).  Punta  Hughes,  SIO  64-45,  1(109). 

Gulf  of  California:  La  Paz,  SIO  74-81,  1(200).  Punta  Man- 
gles, SIO  65-335,  1(122).  Isla  Santa  Inez,  SIO  65-306,  1(99). 
San  Felipe,  SIO  67-1,  1(168). 

Mazatlan  South:  Boca  Teacapan,  SIO  60-90,  1(118).  Nay- 
arit,  SIO  60-89,  1(92).  Bahfa  de  Banderas,  SIO  62-29,  1(148). 
Acapulco,  UCLA  W52-119,  1(70).  Golfo  de  Tehuantepec,  SIO 


73-258,  5(1 16-136);  SIO  68-16,  1(47);  SIO  63-1031,  1(115);  SIO 
72-123,  1(80).  Guatemala,  SIO  63-623,  1(70);  UCLA  W56-273, 
1(120).  Costa  Rica,  UCLA  W54-139,  1(114).  Panama,  SIO  71- 
260,  1(115);  SIO  71-186,  1(36):  USNM  82063,  1(66),  holotype 
of  H.  hildebrandi;  USNM  82037,  1(44),  paratype  of  H.  hil- 
debrandi: USNM  82039,  1(47),  paratype  of  H.  hildebrandi. 
Galapagos  Islands:  BC  56-440,  2(238-247);  SIO  54-174, 
1(128).  Isla  Santa  Cruz,  SIO  H5 1-388,  1(89). 

Doryrhamphus  Kaup 

Doryrhamphus  KAUP,  1853:233  (nomen  nudum);  1856:54 
(type-species  by  monotypy,  D.  excisus  Kaup:  Red  Sea); 
DUMERIL  1870:585  (description);  JORDAN  AND  EVERMANN 
1896:773  (in  part:  description);  DUNCKER  1912:231  (descrip- 
tion; synonymy);  1915:244  (description;  synonymy);  JOR- 
DAN, EVERMANN,  AND  CLARK  1930:243  (synonymy):  HER- 
ALD 1953:244  (description;  synonymy). 

Pristidoryrhamphus  FOWLER,  1944:158  (type-species  by  orig- 
inal designation,  P.  jacksoni  Fowler  =  Doryrhamphus  ne- 
grosensis  Herre). 

DIAGNOSIS. — Gastrophori  with  lateral  trunk 
ridge  continuous  with  inferior  tail  ridge;  inferior 
trunk  and  tail  ridges  discontinuous;  superior 
trunk  and  tail  ridges  discontinuous;  trunk  rings 
more  numerous  than  tail  rings;  snout  ridge 
strongly  serrate,  markedly  so  in  mature  males; 
each  dermal  plate  armed  with  strong  retrorse 
spine;  no  pouch-protecting  plates;  caudal  fin 
large  and  brightly  colored;  branchial  skeleton 
with  all  elements  present;  scutella  present;  two 
nuchal  plates;  three  infraorbitals;  strongly  de- 
veloped opercular  ridge;  14-19  trunk  rings;  10- 
17  tail  rings;  21-29  dorsal  rays;  4  anal  rays;  10 
caudal  rays. 

RANGE. — Four  or  five  species  ranging 
throughout  the  tropical  Indo-Pacific  among  coral 
and  rocky  reefs. 

DISCUSSION. — Kaup  first  published  the  name 
Doryrhamphus  in  1853,  but  his  reference  to  D. 
excisus  Hemprich  and  Ehrenberg  cannot  be  tak- 
en to  be  an  indication  as  defined  by  the  Inter- 
national Code  of  Zoological  Nomenclature  Art. 
16a(v).  The  species  D.  excisus  was  an  unpub- 
lished manuscript  name  in  1853.  The  require- 
ments of  the  Code  were  not  fulfilled  until  Kaup 
(1856)  published  descriptions  of  Doryrhamphus 
and  D.  excisus. 

Key  to  Eastern  Pacific  Species  of 
Doryrhamphus 

la.  Trunk  rings  16-18  (usually  17)  tail  rings 
14-17  (usually  15);  head  4.0-4.4  in  SL  _ 
melanopleura  (Bleeker) 

Tropical  Indo-Pacific 

lb.  Trunk  rings  16-17  (usually  16);  tail  rings 


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' 


FIGURE  4.  Anterior  part  of  body  of  four  species  of  eastern  Pacific  Syngnathidae.  (A)  Leptonotus  blainvilleanus,  a  168-mm- 
SL  female,  SIO  72-168.  (B)  Doryrhamphus  paulus,  holotype,  SIO  72-67.  (C)  Doryrhamphus  melanopleura,  a  52-mm-SL  male, 
SIO  65-343.  (D)  Syngnathus  auliscus,  a  110-mm-SL  female,  SIO  65-181. 


13-15  (usually  14);  head  3.4-4.0  in  SL  _ 
paulus  n.sp. 

Islas  Revillagigedo,  Mexico 

Doryrhamphus  melanopleura  (Bleeker) 

(Figure  4C) 

Syngnathus  melanopleura   BLEEKER,    1858:464  (original  de- 
scription; "Kokos-eilanden"). 


Doryrhamphus  californiensis  GILL,  1863:284  (original  descrip- 
tion; Cabo  San  Lucas;  holotype  SU  19255);  JORDAN  AND 
EVERMANN  1896:773  (description:  range);  DUNCKER  1915:62 
(as  "species  dubia");  ULREY  1929:  6  (Cape  San  Lucas); 
JORDAN,  EVERMANN,  AND  CLARK  1930:243  (range):  ULREY 
1932:77  (Cabo  San  Lucas);  NICHOLS  AND  MURPHY  1944:239 
(Panama);  FOWLER  1944:496  (range). 

Doryichthys  californiensis:  GUNTHER  1870:186  (new  combi- 
nation; description;  range). 


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189 


Microphis  extensus  SNYDER  1911:525  (original  description; 
Naha,  Okinawa;  holotype  USNM  98266). 

Doryrhamphus  melanopleura:  HERALD  1940:59  (in  key;  syn- 
onymy; range);  1953:246  (description;  subspecies;  range). 

DIAGNOSIS. — A  Doryrhamphus  with  22-27 
(24-25)  dorsal  rays;  19-23  pectoral  rays;  16-18 
(17)  trunk  rings;  14-17  (15)  tail  rings;  dorsal  cov- 
ering 3-5  +  2-4  rings;  total  rings  covered  by 
dorsal  fin  6-7.5;  brood  pouch  covering  17-19 
rings;  head  4.0-4.4  in  SL;  dorsal  fin  base  1.6- 
1.9  in  head;  snout  2.0-2.5  in  head;  maximum 
known  size  69  mm  SL;  smallest  mature  male  31 
mm  SL. 

DESCRIPTION. — A  sharply  ridged  and  strongly 
spined  fish.  Head  with  prominent  prenuchal,  nu- 
chal, and  occipital  crests.  Snout  ridge  strongly 
serrated,  markedly  so  in  mature  males.  Pectoral 
cover  plate  with  well-developed  superior  and  in- 
ferior ridges.  Strong  spination  on  maxillae  and 
along  dorsal  edge  of  infraorbitals.  Ventral  edge 
of  quadrate  occasionally  with  strong  serrations. 
Each  ring  bears  a  single  posteriorly  directed 
spine  on  each  ridge,  becoming  obsolete  along 
ventral  ridges.  Scutella  large.  No  dermal  flaps. 

Brood  pouch  abdominal,  without  protecting 
plates.  Eggs  about  0.5  mm  in  greatest  diameter, 
arranged  in  three  layers,  four  across.  Males  with 
eggs  and  young  collected  from  March  through 
August. 

Fins  all  large  and  well  developed.  Caudal  a 
little  longer  than  snout.  Dorsal  base  about  equal 
to  snout  and  orbit  combined.  Anal  conspicuous, 
its  length  about  half  orbital  diameter.  Pectoral 
fins  broad  based. 

Coloration  in  life.  Essentially  the  same  as  col- 
oration in  alcohol  except  that  caudal  has  a  white 
border,  two  yellowish  spots  near  base,  and  an 
oval  orange  area  in  middle  part  of  fin. 

Coloration  in  alcohol.  Uniform  dusky  with 
darker  streak  from  tip  of  snout  to  pectoral  base. 
Fins  colorless  except  for  caudal,  which  is  dis- 
tinctively marked  with  two  light  basal  spots  and 
one  larger  median  spot.  Individual  fin  rays  often 
lined  with  melanophores  on  each  side.  Young 
less  than  20  mm  SL  have  a  banded  color  pattern 
with  eight  brown  bands  on  a  lighter  background. 

HABITAT. — In  crevices  in  rocks,  under  over- 
hangs, and  among  coral  heads. 

RANGE. — In  eastern  Pacific  from  Bahia  Mag- 
dalena,  Baja  California,  south  to  Isla  la  Plata, 
Ecuador  (UMML),  and  at  the  Galapagos  Islands 
and  Clipperton  Island.  In  the  Gulf  of  California 
north  to  Isla  Angel  de  la  Guarda  (UCLA  W60- 


6,  not  examined)  in  the  west  to  Punta  Guillermo, 
Sonora  (UCLA  W51-11  not  examined),  in  the 
east.  Widely  distributed  throughout  Indo-Pacif- 
ic. 

COMPARISONS. — D.  melanopleura  is  an  inter- 
esting and  complex  species.  It  has  been  divided 
into  several  subspecies  (Herald  1953)  that  are 
probably  not  valid.  A  detailed  study  of  the  pop- 
ulations is  needed  before  much  can  be  said  about 
possible  relationships.  It  is  related  to  D.  paulus 
from  which  it  can  be  distinguished  by  the  char- 
acters given  in  the  description  of  that  species. 

MATERIAL  EXAMINED. — Mexico:  Bahia  Magdalena,  SIO  64- 
54,  3(55-62).  Cabo  San  Lucas,  SIO  61-227,  10(18-56).  Los 
Frailes,  SIO  61-249,  14(32-64).  Bahia  de  !os  Muertos,  SIO  74- 
90,  3(32-46).  Isla  San  Jose,  SIO  65-265,  5(48-55).  Isla  Santa 
Cruz,  SIO  65-343,  6(19-61).  Isla  Carmen,  SIO  65-302,  15(45- 
62).  Isla  San  Ignacio  de  Farallon,  SIO  59-228,  3(40-42).  Lobos 
Rock,  SIO  61-280,  20(16-53).  Bahia  de  Banderas,  SIO  62-29, 
2(29-45). 

Costa  Rica:  Isla  del  Cano,  LACM  32548,  36(33-55). 

Panama:  Taboguilla  Island,  SIO  67-34,  21(16-48). 

Galapagos  Islands:  Plaza  Island,  SIO  64-1015,  3(53-63);  BC 
54-392,  1(64). 

Comparative  material  from  Indo-West  Pacific:  Hawaii,  CAS 
20402,  1(54).  Eniwetak,  R.  S.  Nolan  personal  collection,  3(39- 
45).  Guam,  CAS  15835,  2(44-44).  Australia,  AMS  IA-2713, 
1(38);  AMS  110740,  2(31-39).  Japan,  CAS  14512,  2(68-69). 

Doryrhamphus  paulus  n.sp. 
(Figure  4B) 

Doryrhamphus  melanopleura  pleurotaenia:  (in  part)  HERALD 

1953:248  (size;  meristic  data). 
Doryrhamphus  melanopleura:  Ricker  1959:2  (Socorro  Island). 

DIAGNOSIS. — A  dwarf  species  of  Doryrham- 
phus with  23-26  dorsal  rays;  20-21  pectoral 
rays;  16-17  (16)  trunk  rings;  13-15  (14)  tail  rings; 
29-31  (30)  total  rings;  dorsal  on  3-4.2  +  2.5^t 
rings;  total  rings  covered  by  dorsal  fin  6.5-7.5; 
brood  pouch  on  14-17  rings;  head  3.4-4.0  in  SL; 
dorsal  fin  base  1.7-2.2  in  head;  snout  2.1-2.7  in 
head;  smallest  mature  male  24.5  mm  SL;  maxi- 
mum known  size  38.9  mm  SL. 

DESCRIPTION. — Strongly  ridged  and  spined. 
Occipital,  nuchal,  and  prenuchal  ridges  low  but 
sharply  defined.  Snout  ridge  serrate  in  males, 
with  4-5  spines  on  anterior  tip  and  three  or  so 
larger  isolated  spines  further  posterior;  females 
and  juveniles  with  or  without  obvious  snout- 
ridge  serrations.  Pectoral  cover  plate  with  su- 
perior and  inferior  ridges.  Each  body  plate  with 
a  sharp  retrorse  spine.  No  dermal  flaps. 

Only  one  brooding  male  known,  collected  in 
March. 

Fins  well  developed.  Caudal  shorter  than 


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snout.  Dorsal  fin  base  equal  to  combined  snout 
and  orbit.  Pectoral  fins  broad  based  but  short, 
only  extending  posteriorly  to  junction  of  first 
and  second  trunk  rings. 

Coloration.  Essentially  same  as  that  given  for 
eastern  Pacific  populations  of  D.  melanopleura. 

HABITAT  AND  RANGE. — Three  to  17  m, 
among  rock  reefs  in  Islas  Revillagigedo,  Baja 
California. 

COMPARISONS. — D.  paulus  is  most  closely  re- 
lated to,  and  was  probably  derived  from,  D, 
melanopleura,  which  is  widespread  in  the  Indo- 
Pacific  but  is  not  represented  at  the  Islas  Revil- 
lagigedo. Although  D.  melanopleura  ranges 
throughout  the  Indo-Pacific  and  is  quite  vari- 
able, the  number  of  trunk  rings  is  fairly  stable 
at  18.  Some  populations  have  modal  counts  of 
17  or  19  rings;  however,  16  trunk  rings  have  nev- 
er been  found  in  D.  melanopleura. 

Doryrhamphus  paulus  can  be  distinguished 
from  D.  melanopleura  by  the  characters  given 
in  the  key. 

D.  excisus  from  the  Red  Sea  also  has  16  trunk 
rings,  but  differs  from  D.  paulus  in  having  10- 
11  tail  rings  rather  than  13-15,  and  18-20  dorsal 
rays  rather  than  23-26. 

ETYMOLOGY. — From  the  Latin  paulus,  little. 

MATERIAL  EXAMINED.— Holotype:  SIO  72-67,  a  32.7-mm- 
SL  mature  male  collected  at  a  depth  of  6-12  m  with  Chemfish, 
ca.  100  m  SE  of  "Humpback  Cove,"  Isla  Socorro,  Islas  Re- 
villagigedo, Mexico,  19  Feb.  1972,  by  D.  Diener  and  party. 

Paratypes.  Mexico.  Islas  Revillagigedo:  SU  67255,  24.5  mm 
SL,  mature  male,  8  m,  rotenone,  "Grayson's  Cove,"  Isla  So- 
corro, 11  Mar.  1940,  by  Vernon  Brock.  SU  36442,  1(31),  same 
data  as  SU  67255.  CAS  13699,  9(20-34)  and  LACM  31781-2, 
3(22-32),  "Grayson's  Cove,"  ca.  200  m  N  of  "Old  Man  of  the 
Rocks,"  Isla  Socorro,  R/V  SEARCHER  sta.  52,  8-12  m,  rote- 
none, 14  Feb.  1971.  LACM  317821-12,  5(26-32),  s  of  Cape 
Henslow,  Isla  Socorro,  R/V/SEARCHER  sta.  53,  13-17  m,  ro- 
tenone, 15  Feb.  1971.  GCR1>  15753,  1(27),  Braithwaite  Bay, 
Isla  Socorro,  3  m,  rotenone,  13  Feb.  1956,  by  George  Lindsay. 
SIO  72-67,, 6(16-26),  collected  with  holotype.  SIO  74-155, 
2(29-39),  Sulfur  Bay,  Isla  Clarion,  10-17  m,  rotenone,  1 1  Dec. 
1974,  by  Robert  Kiwala. 

Leptonotus  Kaup 

> 

Leptonotus  KAUP,  1853:232  (type-species  by  .monotypy, 
Syngnathus  blainvilleanus  Eydoux  and  Gervais,  1837): 
1856:46;  DUNCKER  1912:235;  1915:88. 

Acmonotus  PHILLIPI,  1896:382  (type-species  by  original  des- 
ignation, Acmonotus  chilensis  .Philippi  [=5.  blainvilleanus 
Eydoux  and  Gervais]). 

Novacampus  WHITLEY,  1955:110  [type-species  by  original 
designation,  Syngnathus  norae  (Waite)]. 

DIAGNOSIS. — Urophori  without  pouch-pro- 
tecting plates;  trunk  compressed  and  much  ex- 


panded in  females;  lateral  ridge  system  of  the 
ascending  pattern  (Fig.  2fi);  all  elements  of  bran- 
chial skeleton  present,  none  reduced;  dorsal  fin 
usually  located  on  two  or  more  trunk  rings;  10 
caudal  rays;  opercular  ridge  weak  or  absent; 
most  body  ridges  smooth  and  reduced;  two 
branchiostegals. 

DISTRIBUTION. — Approximately  five  species 
known  only  from  south  temperate  seas;  South 
America,  South  Australia,  Tasmania,  and  New 
Zealand. 

Leptonotus  blainvilleanus  (Eydoux  and  Gervais) 

(Figure  4A) 

Syngnathus  blainvilleanus  EYDOUX  AND  GERVAIS,  1837:79 
(original  description;  "Mare  Indicum";  holotype  MNHN 
6050);  GUICHENOT  1848:348  (description). 

Syngnathus  acicularis  JENYNS,  1842:147  (original  description; 
Valparaiso,  Chile;  holotype  BMNH  1917.7.14.28). 

Leptonotus  Blainvillei:  KAUP  1853:232  (range);  1856:16  (de- 
scription; range). 

Syngnathus  blainvillianus;  GUNTHER  1870:162  (description: 
range);  STEINDACHNER  1898:331  (Chile):  THOMPSON  1816:423 
(Patagonia). 

Hemithylacus  Petersi  DUMERIL,  1870:600  (original  descrip- 
tion; Puerto  Montt,  Chile). 

Acmonotus  chilensis  PHILIPPI,  1896:382  (original  description; 
Pelluhue,  Chile). 

Leptonotus  blainvillianus:  ABBOTT  1899:338  (references; 
range);  DUNCKER  1915:88  (synonymy;  description;  range); 
NORMAN  1937:40  (Patagonia);  HERALD  1940:59  (synony- 
my; range);  1942:132  (diagnosed  in  key);  FOWLER  1944:4% 
(range);  MANN  1954:189  (description;  range);  DE  BUEN 
1963:89  (synonymy;  description);  KOEPCKE  1962:200  (range); 
CHIRICHIGNO-F.  1974:339  (in  keys;  range). 

Syngnathus  blainvillei:  DELFIN  1901:43. 

Siphostoma  blainvilliana:  EVERMANN  AND  RADCLIFFE 
1917:53. 

Leptonotus  blaenvillianus:  SICCARDI  1954:211-242  (brood 
pouch;  variation). 

Leptonotus  blainvilleanus:  DUMERIL  1870(2):581  (description; 
habitat);  VAILLANT  1888:16  (Orange  Bay,  Patagonia;  col- 
oration); HERALD  1965:364  (common  name). 

DIAGNOSIS. — A  Leptonotus  with  34-41  (35- 
37)  dorsal  rays;  12-14  pectoral  rays;  2-3  anal 
rays;  18-20  (19)  trunk  rings;  48-52  (50)  tail  rings; 
67-70  total  rings;  dorsal  on  0.5-2  +  7-8.2  rings; 
total  rings  covered  by  dorsal  fin  7.3-10;  brood 
pouch  on  10-14  rings;  head  7.0-9.0  in  SL;  dorsal 
fin  base  1.0-1.6  in  head;  snout  1.8-2.2  in  head; 
maximum  known  size  217  mm  SL  (Duncker 
1915);  smallest  mature  male  108  mm  SL.  See 
Table  1  for  meristic  variation. 

DESCRIPTION. — Ridges  on  head  and  body 
mostly  obsolete.  Nuchal  and  prenuchal  ridges 
low  and  smooth.  Opercles  strongly  convex; 
opercular  ridge  reduced  to  basal  one-fourth  of 


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191 


TABLE  1.     MERISTIC  VARIATION  IN  Leptonotus  blainvilleanus. 


Trunk  rings 

Tail  rings 

Dorsal  rays 

Locality                 18 

19 

20 

48        49 

50 

51         52 

34        35        36 

37 

38 

39        40        41 

Chile: 

Antofagasta               1 

12 

1 

1           3 

6 

4 

2          6          1 

4 

1 

—        —        — 

Valparaiso 

— 

2 

—        — 

It 

1        — 

_        —        _ 

It 

— 

_        —        — 

Concepcion 

1 

— 

—        — 

— 

1 

_        _        _ 

1 

— 

—        —        _ 

Coquimbo 

2 

— 

—        — 

1 

1        — 

1         — 

— 

— 

—        —        — 

Puerto  Montt             1 

4 

2 

1           2 

4 

—        — 

1 

3 

3 

_        _        — 

Isla  Chiloe 

9 

3 

4           4 

4 

—        — 

2          3 

4 

2 

—        —        — 

Totals                            2 

28 

8 

6           9 

16 

7        — 

385 

13 

6 

_        _        _ 

x  = 

19.1 

x  =  49.6 

x  = 

36.3 

Argentina*: 

Golfo  Nuevo 

(females)              27 

4 

— 

4 

16 

10          1 

—          1           1 

8 

14 

5          1           1 

(males)                  3 

34 

3 

5          12 

20 

—         2 

7 

7 

13 

8          1           1 

Totals                         30 

38 

3 

5          16 

36 

10         3 

1          8 

15 

27 

13         2          2 

x  = 

18.6 

x  =  49.8 

x  = 

37.8 

"Mare  Indicum"**    — 

1 

— 

—        — 

— 

—        — 

1**     _        _ 

— 

— 

—        —        — 

t  Holotype  of  S.  acicularis. 
*  Data  from  Siccardi  1954. 
**  Holotype  of  L.  blainvilleanus. 


opercle,  striations  faint.  Snout  ridge  smooth, 
low,  and  reaches  posteriorly  to  interorbit.  Su- 
praorbital  ridges  smooth,  one-half  orbit  diameter 
in  length.  Pectoral  cover  plate  without  ridges. 
Trunk  of  mature  females  compressed  and  ex- 
panded dorsoventrally,  with  sharp  dorsal  and 
ventral  borders.  Scutella  small,  indistinct.  Plates 
with  reduced  ridgelets.  Entire  body  very  fleshy 
over  plates.  Lateral  line  papillae  2-4  per  plate. 
No  dermal  flaps. 

Brood  pouch  without  lateral  protecting  plates. 
Males  brooding  eggs  have  been  collected  in  Jan- 
uary and  March. 

Dorsal  fin  high,  its  height  equal  to  width  of 
two  trunk  rings. 

Coloration  in  alcohol.  Juveniles  with  alternat- 
ing pattern  of  dark  and  light  bands  along  length 
of  body.  Darker  bands  about  four  rings  in  width, 
light  bands  only  one-half  a  ring  in  width.  Caudal 
dark  brown  with  a  light  border.  Snout  and  in- 
terorbit darker  than  remainder  of  head.  Fins  col- 
orless. 

Adult  females  sometimes  entirely  light  tan 
with  dark  brown  venter  and  dorsum  on  trunk, 
and  with  posterior  half  of  tail  dark  brown.  Usu- 
ally the  head  is  a  dark  olive-brown  and  the  trunk 
is  dark  brown  with  small,  dark-bordered  ocelli. 


Dorsal  fin  may  have  a  few  melanophores  along 
the  ray  margins.  Adult  males  yellowish  brown 
with  dark  brown  area  surrounding  the  nares; 
ocelli  may  develop  on  the  first  few  tail  rings. 

HABITAT. — Kelp  beds  along  the  open  coast; 
occasionally  into  brackish  water  (Fischer  1963). 

REMARKS. — Meristic  data  from  the  Golfo 
Nuevo,  Argentina,  population  of  L.  blainvil- 
leanus (Table  1)  (from  Siccardi  1954)  show  a 
difference  in  mean  number  of  trunk  rings  be- 
tween males  and  females  from  Golfo  Nuevo  (P 
<  .005). 

The  lower  number  of  trunk  rings  in  the  Golfo 
Nuevo  females  contributes  to  the  significant  dif- 
ference between  the  Chilean  and  Argentinean 
populations  (P  <  .005).  However,  the  dorsal  ray 
count  also  differs  significantly  between  these 
two  regions. 

Siccardi  (1954)  studied  a  large  number  of 
specimens  of  L.  blainvilleanus  from  both  coasts 
of  southern  South  America  and  was  able  to  pro- 
vide some  data  on  morphometric  variation.  She 
found  that  the  depth  of  the  trunk  in  males  re- 
mains fairly  constant  during  growth,  from  112- 
140  mm,  so  that  the  standard  length/trunk  depth 
ratio  was  22.4  in  112-mm  fish  and  26.7  in  140- 
mm  fish.  Also,  the  maximum  depth  of  the  trunk 


192 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


in  females  was  attained  by  a  standard  length  of 
140-150  mm.  This  was  correlated  with  reaching 
sexual  maturity.  The  trunk  in  males  became  rel- 
atively shorter  and  the  tail  longer  with  growth, 
whereas  relative  lengths  of  the  trunk  and  tail  in 
females  remained  constant  with  growth. 

COMPARISONS. — Because  the  genus  Lepton- 
otus  has  not  received  systematic  treatment,  it 
can  only  be  said  that  L.  blainvilleanus  seems  to 
be  most  closely  related  to  the  southern  Austra- 
lian species  L.  semistriatus.  L.  blainvilleanus 
can  be  distinguished  from  L.  semistriatus  by  the 
snout  length  (1.8-2.2  in  head  rather  than  1.6- 
1.7)  and  by  the  dorsal  fin  placement  (on  0.5-2 
+  7-8.2  rings  rather  than  3-4  +  7). 

Leptonotus  blainvilleanus  has  long  been  con- 
sidered to  be  a  member  of  the  New  Zealand  fau- 
na (Waite  1909).  However,  a  comparison  of  the 
holotype  of  L.  blainvilleanus  with  specimens  of 
Leptonotus  from  New  Zealand  reveals  that  L. 
blainvilleanus  and  examined  New  Zealand 
species  of  Leptonotus  are  distinct.  The  New 
Zealand  specimens  of  Leptonotus  are  referable 
to  L.  elevatus  (Hutton)  and  L.  norae  (Waite). 

Mr.  A.  Wheeler  (BMNH)  examined  the  ho- 
lotype of  Syngnathus  acicularis  Jenyns  at  my 
request.  His  counts  and  description  of  lateral 
ridge  pattern  confirm  the  fact  that  the  holotype 
of  S.  acicularis  is  conspecific  with  L.  blainvil- 
leanus. 

RANGE. — Hornitos,  Chile,  to  Golfo  Nuevo, 
Argentina. 

MATERIAL  EXAMINED. — "Mare  Indicum,"  MNHN  6050, 
holotype. 

Chile:  Hornitos,  SIO  72-168,  1(168).  Antofagasta,  EBMCh 
1520-28,  9(86-207);  GCRL  12466,  4(117-215).  Coquimbo, 
MNMHP  5302,  1(169);  MNMHP  5305,  1(246);  Valparaiso, 
EBMCh  222,  1(190).  Concepcion,  MNMHP  5574,  1(126).  Ba- 
hi'a  Lin,  USNM  176564,  4(75-121).  Puerto  Montt,  USNM 
205179,  3(104-129).  Isla  de  Chiloe,  USNM  176569,  4(108- 
142);  MNMHP  5304,  3(84-173);  MNMHP  5303,  5(141-162). 
No  collection  data,  CAS  (Indiana  label),  1(184). 

Argentina:  Patagonia,  CAS  36440,  1(130). 

Bryx  Herald 

Bryx  HERALD,  1940:52  (type-species  by  original  designation 
Bryx  veleronis  Herald);  1959:468  (subgenus  of  Syngnathus). 

Microsyngnathus  HERALD,  1959:468  (subgenus  of  Syngna- 
thus, type-species  by  original  designation  Syngnathus 
dunckeri  Metzelaar). 

DIAGNOSIS. — Urophori  characterized  by  a 
very  short  snout;  small  size  (generally  less  than 
100  mm);  first  epibranchial  reduced  or  not;  sec- 
ond hypobranchials  and  epibranchials  reduced; 


frontals  not  reaching  anteriorly  past  middle  of 
lateral  ethmoids;  ossified  epaxialis  tendons  pres- 
ent or  absent;  one  infraorbital;  two  nuchal 
plates;  pouch  protecting  plates  present;  dermal 
flaps  present  or  absent;  14-17  trunk  rings;  27-40 
tail  rings;  19-31  dorsal  rays;  10-14  pectoral  rays; 
anal  fin  present  or  absent;  10  caudal  rays;  10-19 
rings  covered  by  brood  pouch;  0-2  +  3-7  rings 
covered  by  dorsal  fin. 

DISTRIBUTION. — Ten  tropical  species,  and 
one  species  at  Islas  Juan  Fernandez  and  Isla  San 
Felix. 

DISCUSSION. — Bryx  is  closely  related  to  Cos- 
mocampus,  and  the  two  were  probably  derived 
from  a  common  ancestor. 

Bryx  was  erected  by  Herald  in  1940  for  the 
sole  reception  of  B.  veleronis.  Later,  Herald 
(1959)  transferred  B.  veleronis  to  Syngnathus 
and  retained  Bryx  as  a  subgenus.  In  the  same 
paper  Microsyngnathus  was  erected  as  a  sub- 
genus,  with  Syngnathus  dunckeri  as  type-spe- 
cies, and  S.  arctus  and  S.  coccineus  included 
therein.  S.  hildebrandi  Herald,  1965,  S.  ran- 
dalli  Herald,  1965,  and  5.  banneri  Herald  and 
Randall,  1972,  were  described  as  being  related 
to,  or  tentatively  referred  to,  the  subgenus  Mi- 
crosyngnathus species  in  Herald  (1965)  and  Her- 
ald and  Randall,  1972.  The  species  of  Bryx  and 
Microsyngnathus  (and  Micrognathus  balli  Fow- 
ler, 1925,  and  Syngnathus  darrosanus  Dawson 
and  Randall,  1975)  are  separated  from  other 
syngnathids  and  united  by  shared  characters 
given  in  the  diagnosis  above.  This  combination 
of  characters  defines  a  related  lineage  of  syng- 
nathids worthy  of  generic  recognition.  The  type- 
species  of  Microsyngnathus  further  agrees  with 
Bryx  veleronis  in  lacking  an  anal  fin.  Micro- 
syngnathus Herald,  1959,  thus  becomes  a  junior 
synonym  of  Bryx  Herald,  1940.  The  subgenus 
Simocampus  is  proposed  for  those  species  of 
Bryx  possessing  an  anal  fin. 

Key  to  Eastern  Pacific  Species  of  Bryx 
la.  Anal  fin  present  (subgenus  Simocampus) 
2 

Ib.  Anal  fin  absent  (subgenus  Bryx)  4 

2a.  Head  shorter  than  length  of  dorsal  fin 
base heraldi  n.sp. 

Islas  Juan  Fernandez  and  Isla 
San  Felix 

2b.  Head  longer  than  length  of  dorsal  fin 
base 3 

3a.  Dorsal  fin  usually  entirely  on  tail  rings, 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


193 


TABLE  2.     FREQUENCY  DISTRIBUTIONS  OF  TRUNK  AND  TAIL  RINGS,  AND  DORSAL  FIN  RAYS  IN  EASTERN  PACIFIC  SPECIES 
OF  Brvx. 


Trunk 

rings 

Tail  rings 

Dorsal  fin  rays 

14 

15 

16     33     34     35    36 

37 

38 

39 

40    41     18     19      20 

21     22 

23 

24     25     26     27       28 

heraldi 



18* 

_    _    _    _    2 

10 

6* 



.  

-     10* 

8 



arctus 

1 

31* 

20    -                 -     3 

5 

10 

18* 

19       1       4     19       10* 

10      9 

2 

—    —    —    —      — 

coccineus 

1 

11* 

1     -                  -     1 

3 

2 

5* 

2    -                 8*       4 

—    — 

— 

_    _    _    _      _ 

veleronis 

1 

16* 

2    -         19* 

5 

— 

— 

—    —    —    —      — 

1 

1 

13442 

clarionensis 

— 

9* 

—    —    —      36* 

— 

— 

— 

—    —    —    —      — 

—    — 

— 

1       2       4*       2 

*  Primary  type. 


except  in  some  southern  California  spec- 
imens with  the  dorsal  on  a  fraction  of  a 
trunk  ring;  18-23  dorsal  rays;  double  row 

of  dark  spots  on  trunk  

arctus  (Jenkins  and  Evermann) 

Tomales  Bay,  California,  and 
San  Felipe,  Gulf  of  California, 
to  Mazatlan,  Mexico 

3b.  Dorsal  fin  always  on  at  least  a  fraction  of 
last  trunk  ring;  18-20  dorsal  rays;  no  dou- 
ble row  of  dark  spots  on  trunk 

coccineus  (Herald) 

Banderas  Bay,  Mexico,  to 
Punta  Aguja,  Peru,  and  Gala- 
pagos Islands 

4a.  Snout  longer  than  postorbital  length  

clarionensis  n.sp. 

Clarion  Island,  Revillagigedo 
Islands,  Mexico 

4b.  Snout  shorter  than  postorbital  length  .... 
veleronis  Herald 

Galapagos  and  Revillagigedo 
Islands,  and  coasts  of  Panama 
and  Costa  Rica 

Simocampus  n.subgen. 

TYPE-SPECIES.- — Siphostoma  arctum  Jenkins  and  Evermann. 

DIAGNOSIS. — A  Bryx  with  anal  fin. 


INCLUDED  SPECIES. — B.  arctus  (Jenkins  and 
Evermann);  B.  coccineus  (Herald);  B.  balli 
(Fowler);  B.  banneri  (Herald  and  Randall);  B. 
hildebrandi  (Herald);  and  B.  darrosanus  (Daw- 
son  and  Randall). 

ETYMOLOGY. — From  the  Greek  simos,  pug- 
nosed,  and  campos,  sea-creature. 

Bryx  (Simocampus)  heraldi  n.sp. 

(Figure  5C) 

DIAGNOSIS.— A  Bryx  with  22-23  (22)  dorsal 
rays;  11  pectoral  rays;  3  anal  rays;  15  trunk 
rings;  36-38  (37)  tail  rings;  52-54  total  rings;  dor- 
sal on  0-1  +  5.2-6  rings;  17-18  rings  covered 
by  brood  pouch;  head  10.3-12.3  in  SL;  dorsal 
fin  base  0.8-1.0  in  head;  snout  2.4-2.8  in  head; 
maximum  known  size  99.8  mm  SL;  smallest 
mature  male  70  mm  SL.  See  Tables  2  and  3  for 
additional  counts  and  measurements. 

DESCRIPTION. — All  ridges  of  head  and  body 
moderately  developed  and  smooth.  Prenuchal 
and  nuchal  ridges  elevated  and  easily  discern- 
ible. Opercular  ridge  extends  posteriorly  about 
half  length  of  opercle.  Snout  ridge  moderately 
developed,  extending  posteriorly  to  interorbit. 
Ridges  of  pectoral  cover  plate  obsolete.  Trunk 
and  tail  ridges  low  but  easily  visible.  Superior 


TABLE  3.     NUMBER  OF  SPECIMENS  (N)  AND  STANDARD  LENGTH  RANGE,  TOGETHER  WITH  RANGE  AND  MEAN  (?)  OF  SE- 
LECTED CHARACTERS  EXPRESSED  IN  THOUSANDTHS  OF  STANDARD  LENGTH  IN  Bryx. 


Head 

Snout 

Snout 

Body 

Dorsal  base 

Pectoral 

length 

length 

depth 

depth 

length 

fin  length 

Species 

SL 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

N 

heraldi 

70-100 

81-97 

89 

29-37 

33 

12-16 

14 

29-38 

34 

91-113 

100 

16-21 

18 

7 

coccineus 

50-116 

90-115 

100 

35-42 

38 

12-22 

16 

30-47 

36 

72-82 

77 

12-22 

18 

7 

arctus 

62-88 

84-99 

92 

23-41 

33 

12-17 

15 

35-48 

41 

80-87 

83 

11-23 

17 

11 

veleronis 

37-49 

96-113 

104 

41-44 

43 

16-19 

17 

35^»4 

38 

108-130 

123 

17-24 

20 

7 

clarionensis 

38-50 

119-131 

125 

39-44 

42 

15-20 

17 

36-45 

40 

121-136 

126 

23-28 

25 

8 

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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


B 


FIGURE  5.     Anterior  part  of  body  of  three  species  of  eastern  Pacific  Syngnathidae.  (A)  Bryx  arctus,  an  88-mm-SL  female, 
SIO  H46-65.  (B)  Bryx  coccineus,  a  116-mm-SL  female,  USNM  220970.  (C)  Bryx  heraldi,  holotype,  SIO  65-626. 


tail  ridge  and  lateral  trunk  ridge  overlap  on  one 
plate.  Scutella  indistinct  and  about  one-fourth  as 
deep  as  adjoining  plates.  Lateral  line  consisting 
of  2-4  papillae  per  plate.  No  dermal  flaps. 

Protecting  plates  of  brood  pouch  weakly  de- 
veloped; pouch  folds  lightly  scalloped.  Eggs  0.5 
mm  in  greatest  diameter,  arranged  in  two  layers 
four  across.  Males  with  eggs  and  young  have 
been  taken  in  early  to  mid-December. 

Caudal  fin  short,  as  long  as  last  two  tail  rings. 

Coloration  in  alcohol.  Background  a  light  tan 
with  few  to  many  dark  blotches  on  body,  which, 
in  some  individuals,  may  make  the  fish  appear 
darker  brown.  The  dark  blotches  may  be  lighter 
centrally,  and  they  may  appear  as  dark  bands  2- 
4  rings  wide,  or  as  thin  bands  between  light  tan 
areas.  Brown  freckling  may  also  occur  dorsally. 
All  fins  are  colorless.  The  head  coloration  is  es- 
sentially like  that  of  the  rest  of  the  body.  Pig- 
mented  part  of  cornea  a  darkish  tan  ring  with 
radially  arranged  light  blotches. 


HABITAT. — Rocky  reefs  and  sandy  bottoms  at 
depths  of  6-23  m. 

COMPARISONS. — B.  heraldi  is  most  closely  re- 
lated to  B.  coccineus  and  B.  arctus  of  the  east- 
ern Pacific.  It  differs  from  coccineus  is  having 
a  shorter  head  (10-13  rather  than  8.6-11.1  in 
SL),  22-23  rather  than  19-20  dorsal  rays,  and  a 
longer  dorsal  fin  base  (0.8-1.0  rather  than  1.1- 
1.4  in  head  length);  and  from  arctus  in  lacking 
the  double  row  of  dark  spots  on  the  trunk  and 
dermal  flaps,  and  in  having  fewer  tail  rings  (36- 
38  rather  than  36-41). 

RANGE. — Islas  Juan  Fernandez  and  Isla  San 
Felix. 

ETYMOLOGY. — Named  in  honor  of  the  late 
Earl  S.  Herald  in  recognition  of  his  life-long  in- 
terest in  pipefish. 

MATERIAL  EXAMINED.— Holotype:  SIO  65-626,  a  99.8-mm- 
SL  mature  male  collected  from  a  rock  ledge  at  Isla  San  Felix, 
Chile,  from  a  depth  of  9-12  m,  with  emulsified  rotenone  and 
SCUBA  on  6  Dec.  1965  by  Wayne  Baldwin  and  party. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


195 


Paratypes:  SIO  65-626,  2(79-82)  taken  with  the  holotype. 
SIO  65-624,  1(85);  Isla  San  Felix,  9  m.  SIO  65-634,  1(77); 
Cumberland  Bay,  Isla  Juan  Fernandez,  6-1 1  m.  GCRL  15755, 
1(70);  Isla  Juan  Fernandez,  20-23  m.  SIO  65-655,  1(72);  Car- 
vajal  Bay,  Isla  Juan  Fernandez,  9-12  m.  All  taken  5-15  Dec. 
1965  with  emulsified  rotenone  and  SCUBA  by  Wayne  Baldwin 
and  party. 

Bryx  (Simocampus)  arctus  (Jenkins  and  Ever- 
mann) 

(Figures  5A,6,  16B) 

Siphostoma  arctum  JENKINS  AND  EVERMANN,  1888:137  (orig- 
inal description;  Guaymas,  Sonora,  Mexico;  holotype 
USNM  39369);  JORDAN  AND  EVERMANN  1896:771  (descrip- 
tion). 

Syngnathus  arctus:  ULREY  AND  GREELEY  1928:42  (synony- 
my; range);  ULREY  1929:6  (Lower  California;  Gulf  of  Cali- 
fornia); JORDAN,  EVERMANN,  AND  CLARK  1930: 242  (range): 
STARKS  AND  MORRIS  1907:185  (range);  HUBBS  1916:160 
(eelgrass;  San  Diego,  California):  ULREY  1932:77  ("costa 
occidental  y  Golfo  de  California");  HERALD  1940:60  (syn- 
onymy); 1959:468  (placed  in  subgenus  Microsyngnathus); 
FEDER,  TURNER,  AND  LIMBAUGH  1974:128  (in  coralline  al- 
gae under  kelp  beds);  MILLER  AND  LEA  1972:89  (descrip- 
tion; range). 

Syngnathus  arcta:  BERRY  AND  PERKINS  1965:668  (larvae  in 
California  Current). 

DIAGNOSIS. — A  Simocampus  with  18-23  (19) 
dorsal  rays;  3  anal  rays;  9-11  pectoral  rays;  14- 
16  (15)  trunk  rings;  36-41  (39-40)  tail  rings;  dor- 
sal on  0-0.8  +  5-5.3  rings;  17-23  rings  covered 
by  brood  pouch;  head  10.1-13.2  in  SL;  dorsal 
fin  base  1.0-1.2  in  head;  snout  2.4—4.0  in  head; 
maximum  known  size  121  mm  SL;  smallest  sex- 
ually mature  male  59  mm  SL.  See  Tables  2  and 
3  for  additional  counts  and  measurements. 

DESCRIPTION. — Ridges  of  trunk  and  tail  dis- 
tinct and  smooth.  Prenuchal  and  nuchal  ridges 
smooth,  not  separated  by  an  indentation.  Oper- 
cular  ridge  distinct  and  about  half  length  of  oper- 
cle.  Snout  ridge  elevated  and  smooth  from  in- 
terorbital  to  tip  of  vomer.  Supraorbital  ridge 
extends  posteriorly  to  center  of  opercle.  Pector- 
al cover-plate  ridges  joined  to  form  an  anteriorly 
directed  semicircle.  Dermal  flaps  often  present 
at  junction  of  plates  along  ridges. 

Protecting  plates  of  brood  pouch  moderately 
developed;  edges  of  pouch  folds  slightly  scal- 
loped. Eggs  0.5  mm  in  greatest  diameter,  ar- 
ranged in  two  layers  four  across.  Males  with 
eggs  and  young  taken  from  May  to  July. 

Coloration  in  alcohol.  Variable  light  tan  to 
dark  brown.  May  be  uniformly  colored  or  var- 
iously mottled  with  light  and  dark.  Usually  a 
double  row  of  dark  spots  on  sides  of  trunk. 
When  patterned,  background  color  is  usually 


CC 


SC       BC       MB      LP      SF 
LOCALITY 


B 


N        6- 


3  ' 


68      70      72       74      76 
TOTAL 


78       80 


FIGURE  6.  Clinal  variation  of  Bryx  (Simocampus)  arctus. 
(A)  Mean  number  of  dorsal  rays  plus  total  rings  plotted  against 
locality.  CC  =  central  California:  SC  =  southern  California: 
BC  =  Baja  California  south  of  Bahia  Sebastian  Vizcaino; 
MB  =  Bahia  Magdalena;  LP  =  La  Paz  to  Bahia  de  Los  An- 
geles; SF  =  San  Felipe.  (B)  Number  of  dorsal  rays  plus  total 
rings:  northern  Gulf  of  California  specimens  included  with 
southern  specimens.  (C)  Number  of  dorsal  rays  plus  total 
rings:  northern  Gulf  of  California  specimens  included  with 
southern  specimens.  Dotted  line  indicates  specimens  from  the 
north:  solid  line  indicates  specimens  from  the  south. 


light,  with  dark  bands  on  about  every  fourth 
ring.  Northern  fish  usually  darker  and  not  mot- 
tled (Starks  and  Morris  1907). 

HABITAT. — Shallow  water,  0-10  m,  in  bays  in 
eelgrass,  and  on  reefs  in  clumps  of  algae,  often 
coralline. 

VARIATION. — Meristic  features:  There  are 
significant  differences  (P  <l  .001)  in  mean  num- 
bers of  trunk  rings,  tail  rings,  and  dorsal  rays 
between  populations  from  the  Gulf  of  California 


1% 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


and  central  California.  The  differences  are  pro- 
duced by  the  clinal  decrease  in  number  of  rings 
and  dorsal  rays  from  north  to  south  (Fig.  6).  It 
seems  reasonable  to  attribute  this  cline  to  the 
direct  effect  of  the  environmental  temperature. 

COMPARISONS. — B.  arctus  is  most  closely  re- 
lated to  B.  coccineus  and  B.  heraldi  in  the  east- 
ern Pacific  and  B.  hildebrandi  in  the  western 
Atlantic.  B.  arctus  can  be  distinguished  from  B. 
coccineus  and  B.  heraldi  by  the  characters  given 
in  the  key  and  in  the  descriptions  of  these 
species.  B.  arctus  can  be  distinguished  from  B. 
hildebrandi  by  the  lower  number  of  trunk  rings 
(14-16  rather  than  17),  the  higher  number  of  tail 
rings  (36-41  rather  than  33-34),  and  the  smaller 
head  (10.1-13.7  rather  than  9.5-10.7  in  SL)  in 
the  former. 

RANGE. — Tomales  Bay,  California,  south  to 
Mazatlan,  Mexico,  and  throughout  the  Gulf  of 
California. 

MATERIAL  EXAMINED. — California:  Tomales  Bay,  SU 
36445,  1(74);  Elkhorn  Slough,  CAS  45064,  1(84);  SIO  H45-53, 
1(52).  Santa  Monica,  UCLA  W49-112,  1(60).  San  Pedro,  SIO 
74-120,  1(85);  Santa  Catalina  Island,  SIO  53-185,  2(58-79). 
Newport  Bay,  SIO  H45-70,  7(70-84).  San  Clemente,  SIO 
55-73A,  2(90-94).  La  Jolla,  SIO  58-504,  1(73);  SIO  H45-32, 
1(85). 

Outer  Coast  of  Baja  California:  Bahi'a  de  Todos  Santos,  SIO 
62-475,  1(111).  Bahi'a  San  Quintin,  SIO  63-1055,  3(119-121). 
Bahi'a  Playa  Maria,  SIO  52-168,  1(96).  Punta  San  Rosarito, 
SIO  52-162,  3(76-88).  Laguna  San  Ignacio,  SIO  H50-71,  2(59- 
60).  Bahi'a  Santa  Maria,  SIO  62-734,  1(102).  Bahi'a  Magdalena, 
SIO  62-726,  7(54-91);  SIO  58-61,  1(97). 

Gulf  of  California:  Isla  San  Jose,  SIO  65-265,  1(62).  Bahi'a 
San  Carlos,  CAS  48976,  4(71-84).  Isla  Santa  Cruz,  SIO  74- 
124,  1(59):  Isla  Ildefonso,  SIO  65-330,  1(68).  Bahi'a  de  los  An- 
geles, SIO  61-185,  2(90-98).  San  Felipe,  SIO  72-79,  4(66-81). 
Isla  Tiburon,  UCLA  W56-26,  2(60-67). 

Sonora:  Bahi'a,  Bocochibampo,  SIO  70-84,  1(79).  Guaymas, 
USNM  39639,  1(88),  holotype. 

Bryx  (Simocampus)  coccineus  (Herald) 

(Figure  5B) 

Syngnathus  coccineus  HERALD,  1940:57  (original  description; 
Charles  Island,  Galapagos  Islands:  holotype,  USNM 
101688);  FOWLER  1944:496  (Galapagos  Islands). 

Syngnathus  independencia  HILDEBRAND,  1946: 153  (original 
description;  Bahi'a  Independencia,  Peru;  holotype,  USNM 
127853);  KOEPCKE  1962:200  (range);  CHIRICHIGNO-F. 
1974:89,  351  (diagnosed  in  key;  range). 

DIAGNOSIS. — A  Simocampus  with  19-20  (19) 
dorsal  rays;  2-3  anal  rays;  10-11  pectoral  rays; 
14-16  (15)  trunk  rings;  36-40  (39)  tail  rings;  51- 
54  total  rings;  dorsal  on  0-1  +  4.5-5  rings;  14- 
16  rings  covered  by  brood  pouch;  head  8.6-11.1 
in  SL;  dorsal  fin  base  1.1-1.4  in  head;  snout  2.3- 


3.0  in  head;  maximum  known  size  124  mm  SL; 
smallest  sexually  mature  male  78  mm  SL.  See 
Tables  2  and  3  for  additional  counts  and  mea- 
surements. 

DESCRIPTION. — All  ridges  of  trunk  and  tail 
moderately  developed.  Prenuchal,  nuchal,  su- 
praorbital,  and  snout  ridges  well  developed. 
Opercular  ridge  one-half  length  of  opercle  on 
smaller  specimens,  becoming  one-third  length  of 
opercle  with  growth.  Two  horizontal  ridges  on 
pectoral  cover  plate,  becoming  obsolete  with 
growth.  Superior  tail  ridge  and  lateral  trunk 
ridge  overlap  for  0-1  plate.  Lateral  line  with  3- 
6  papillae  per  plate.  Dermal  flaps  may  be  pres- 
ent. 

Protecting  plates  of  brood  pouch  weakly  de- 
veloped and  no  deeper  than  one-half  depth  of 
adjacent  tail;  edges  of  pouch  folds  smooth  to 
weakly  scalloped.  Eggs  0.6  mm  in  greatest  di- 
ameter, arranged  in  one  layer  two  or  three 
across.  Males  with  young  have  been  taken  at  the 
Galapagos  Islands  in  January.  Brooding  males 
have  been  collected  in  July  in  Panama. 

Caudal  fin  as  long  as  last  four  tail  rings. 

Coloration  in  alcohol.  Coloration  in  B.  coc- 
cineus is  very  variable.  Background  color  varies 
from  a  light  tan  to  dark  brown.  Specimens  may 
be  a  uniform  light  tan  to  dark  brown,  or  more 
usually,  light  with  darker  markings.  Light  tan 
specimens  may  have  darker  spots  at  the  bases 
of  the  dorsal  rays  and  pectoral  rays.  The  spec- 
imens with  tan  background  and  darker  blotches 
do  not  have  any  unifying  color  pattern.  One 
specimen  has  the  venter  dark  brown  and  dorsum 
light  tan,  with  dark  bands  extending  over  the 
dorsum  on  every  three  or  four  rings.  Many  spec- 
imens, however,  have  a  whitish  blotch  on  the 
pectoral  cover  plate  and  immediately  posterior 
to  the  orbit.  Hyoid  region  may  be  dark  with  a 
series  of  whitish  spots. 

HABITAT. — Beds  of  red  algae  and  coral  heads 
at  depths  from  0-18  m. 

COMPARISONS. — B.  coccineus  is  most  closely 
related  to  its  eastern  Pacific  congeners  B.  her- 
aldi and  B.  arctus.  B.  coccineus  differs  from  the 
former  by  the  characters  discussed  in  the  de- 
scription of  B.  heraldi,  and  from  the  latter  in 
lacking  the  double  row  of  dark  spots  on  the 
trunk  and  in  having  a  longer  head  (8.6-1 1 . 1  rath- 
er than  10.1-13.2  in  SL).  Examined  type  mate- 
rial of  S.  independencia  agrees  in  all  respects 
with  the  holotype  of  B.  coccineus.  It  is  likely 
that  the  exceptional  occurrence  of  B.  coccineus 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


197 


in  Independencia  Bay,  Peru,  in  1941  was  the  re- 
sult of  transport  of  individuals  southward  during 
the  extreme  El  Nino  of  that  year  (Lobell  1942). 
RANGE. — Bahfa  Banderas,  Mexico,  to  Punta 
Aguja,  Peru  (occasionally  to  Bahfa  Independen- 
cia), and  the  Galpagos  Islands. 

MATERIAL  EXAMINED. — Mexico:  Bahfa  de  Banderas,  SIO 
62-42,  1(116). 

Costa  Rica:  Puerto  Culebra,  USNM  101690,  1(57).  Puerto 
Jimenez,  USNM  92120,  1(50).  Both  paratypes  of  S.  cocci- 
neus. 

Panama:  Isla  Canal  de  Afuera,  SIO  71-52,  1(65). 

Peru:  6°55'S,  80°42'W,  USNM  220970,  1(116).  Bahfa  Inde- 
pendencia, USNM  127853,  1(124),  holotype  of  S.  indepen- 
dencia;  USNM  127854,  2(88-108),  paratypes  of  5.  Indepen- 
dencia. 

Galapagos  Islands:  Isla  Santa  Maria,  USNM  101688,  1(87), 
holotype  of  S.  coccineus.  Isla  Fernandina,  USNM  220971, 
6(62-98):  USNM  220972,  1(77). 

Subgenus  Bryx  Herald 

TYPE-SPECIES. — Bryx  veleronis  Herald. 

DIAGNOSIS. — A  Bryx  without  an  anal  fin. 

INCLUDED  SPECIES. — B.  veleronis  Herald;  B. 
clarionensis  n.sp.;  B.  dunckeri  (Metzelaar);  B. 
randalli  (Herald). 

Bryx  (Bryx)  veleronis  Herald 

(Figures  IB  &  8) 

Bryx  veleronis  HERALD,  1940:55  (original  description;  Albe- 
marle  Island,  Galapagos  Islands;  holotype  LACM  20012). 

DIAGNOSIS. — A  Bryx  with  22-28  dorsal  rays; 
11-13  pectoral  rays;  14-15  trunk  rings;  33-37 
tail  rings;  48-52  total  rings;  dorsal  on  0-0.5  + 
5.5-7.0  rings;  11.5-15  rings  covered  by  brood 
pouch;  head  8.3-10.3  in  SL;  dorsal  fin  base  0.8- 
1.0  in  head;  snout  1.2-1.5  in  postorbital;  maxi- 
mum known  size  60  mm  SL;  smallest  sexually 
mature  male  45  mm  SL.  See  Figure  8,  and  Ta- 
bles 2  and  3  for  additional  counts  and  measure- 
ments. 

DESCRIPTION. — Ridges  of  trunk  and  tail  mod- 
erately developed,  serrate  and  with  an  indenta- 
tion at  plate  junctions.  Prenuchal  and  nuchal 
plates  with  sharply  defined  serrate  ridge.  Snout 
ridge  well  developed,  wholly  on  mesethmoid. 
Opercular  ridge  three-fourths  length  of  opercle. 
Supraorbital  ridge  weak.  Two  parallel  horizontal 
ridges  on  pectoral  cover  plate.  Superior  tail  and 
median  trunk  ridges  overlap  for  one  ring.  Scu- 
tella  indistinct  and  half  as  deep  as  adjoining 
plates.  Lateral  line  consisting  of  1-2  papillae  per 
plate.  Dermal  flaps  may  be  present,  especially 
on  upper  surface  of  head. 


B 


FIGURE  7.  Anterior  part  of  body  of  three  species  of  eastern 
Pacific  Syngnathidae.  (A)  Bryx  clarionensis,  holotype,  SIO 
74-116.  (B)  Bryx  veleronis,  a  34-mm-SL  female,  SIO  67-40. 
(C)  Hippocampus  ingens,  in  life. 


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Protecting  plates  of  brood  pouch  weakly  de- 
veloped and  no  deeper  than  half  depth  of  adja- 
cent plate;  edges  of  pouch  folds  smooth.  Eggs 
0.5-0.75  mm  in  greatest  diameter,  arranged  in 
one  layer  two  across.  Males  with  young  taken 
in  July. 

Coloration  in  alcohol.  Body  a  light  to  dark  tan, 
with  various  amounts  of  mottling,  but  no  definite 
pattern.  Opercular  membrane  and  hyoid  region 
may  be  brownish,  with  white  spots  and  bars  al- 
most serially  arranged.  Pectoral  and  dorsal  fins 
clear,  caudal  colored  the  same  as  rest  of  body. 
Body  may  have  sprinkling  of  punctate  melano- 
phores  ventrally.  One  or  two  brownish  streaks 
may  extend  posteriorly  from  orbit. 

HABITAT. — In  red-algae  beds  and  coral  heads 
at  depths  of  3-18  m. 

COMPARISONS. — Most  closely  related  to  B. 
clarionenesis.  For  further  comparisons  see  dis- 
cussion under  that  species. 

RANGE. — Taken  only  at  islands,  including  the 
Galapagos  Islands;  Islas  Revillagigedo;  Isla 
Murcielago  and  Isla  del  Cano,  Costa  Rica;  and 
Islas  San  Jose  and  Canal  de  Afuera,  Panama. 

MATERIAL  EXAMINED. — Islas  Revillagigedo:  Isla  Clarion, 
LACM  20013  (formerly  AHF  VELERO  sta.  305-34),  13(45-49), 
paratypes.  Isla  Socorro,  UCLA  W55-124,  1(41). 

Costa  Rica:  Isla  del  Cano,  LACM  31579-15,  6(37-48). 

Panama:  Archipielago  de  las  Perlas,  SIO  67-38,  1(37);  SIO 
67-40,  1(34).  Isla  Canal  de  Afuera,  SIO  71-52,  1(46). 

Galapagos  Islands:  Isla  Santa  Maria,  USNM  101689,  1(48), 
paratype. 

Bryx  (Bryx)  clarionensis  n.sp. 

(Figures  7 A  &  8) 

DIAGNOSIS. — A  Bryx  with  25-28  dorsal  rays; 
11-13  pectoral  rays;  15  trunk  rings;  35-36  tail 
rings;  50-51  total  rings;  dorsal  on  0-1  +  6-7 
rings;  head  7.6-8.4  in  SL;  dorsal  fin  base  0.9- 
1.1  in  head;  snout  0.8-0.9  in  postorbital;  maxi- 
mum known  size  47  mm  SL;  males  unknown. 
See  Figure  8,  and  Tables  2  and  3  for  additional 
counts  and  measurements. 

DESCRIPTION. — All  ridges  of  head  and  body 
distinct  and  moderately  serrate.  Prenuchal  and 
nuchal  ridges  slightly  elevated.  Opercular  ridge 
extends  posteriorly  for  entire  length  of  opercle. 
Snout  ridge  extends  from  interorbital  region  to 
anterior  end  of  mesethmoid.  Pectoral  cover 
plate  with  two  obsolete  horizontal  ridges.  Trunk 
and  tail  ridges  not  markedly  indented  between 
rings.  Superior  tail  and  lateral  trunk  ridges  do 
not  overlap.  Scutella  small,  indistinct.  Lateral 


line  not  visible.  Many  cirruslike  dermal  flaps  on 
dorsal  surface  of  head  in  a  few  specimens. 

Coloration  in  alcohol.  Background  color  or- 
angish  tan,  with  a  series  of  light  and  dark  bands 
in  a  few  specimens,  each  about  two  rings  in 
width  around  tail.  Brown  frecklelike  spots  may 
appear  on  dorsal  surface  of  head  and  along  sides 
of  snout.  Abdominal  region  slightly  darker  than 
rest  of  body.  Dorsal  and  pectoral  fins  colorless, 
caudal  rays  orangish  brown. 

HABITAT. — Known  only  from  the  types  taken 
at  the  surface  near  Isla  Clarion,  Mexico. 

COMPARISONS. — B.  clarionenesis  differs  from 
the  closely  related  species  B.  veleronis  in  having 
a  longer  snout  (0.8-0.9  rather  than  1.4-1.5  in 
postorbital)  and  the  snout  ridge  not  quite  so  el- 
evated and  serrate.  Figure  8  provides  a  compar- 
ison of  the  snout  length  versus  standard  length 
for  each  species.  Both  species  have  been  col- 
lected at  Isla  Clarion. 

RANGE. — Isla  Clarion,  Islas  Revillagigedo, 
Mexico. 

ETYMOLOGY. — Named  for  the  type-locality. 

MATERIAL  EXAMINED. — Holotype:  SIO  74-116,  formerly 
UCLA  W55-136,  a  47-mm-SL  mature  female  dip-netted  under 
a  night  light  over  15  fathoms  (27  m)  at  the  SE  anchorage,  Isla 
Clarion,  20  Apr.  1955,  by  John  Fitch,  Wayne  Baldwin,  and  B. 
Craig. 

Paratypes:  SIO  74-116,  7(38-50),  and  GCRL  15754,  1(41), 
same  data  as  holotype. 

Syngnathus  Linnaeus 

Syngnathus  LINNAEUS,  1758:336  (type-species  by  Int.  Comm. 
Zool.  Nom.  Opinions  45  and  77:  Syngnathus  acits  Lin- 
naeus); DUNCKER  1912:236  (description);  1915:78  (descrip- 
tion; subgenera);  JORDAN,  EVERMANN,  AND  CLARK 
1930:240  (North  American  species;  synonymy):  HERALD 
1943:141  (in  part;  description;  subgenera). 

Siphostoma  RAFINESQUE,  1810:18  (type-species  by  mono- 
typy,  Syngnathus  pelagicus  Linnaeus);  JORDAN  AND  EV- 
ERMANN 1896:761  (description;  synonymy). 

Siphonostoma  KAUP,  1853:233  (emmendation  of  Siphosto- 
ma); 1856:48  (description). 

Dermatostethus  GILL,  1863:283  (type-species  by  monotypy, 
Dermatostethus  punctipinnis  Gill  [=Syngnathus  acus  Lin- 
naeus]); JORDAN,  EVERMANN,  AND  CLARK  1930:240 
(range). 

Syrictes  JORDAN  AND  EVERMANN,  1927:504  (type-species  by 
original  designation,  Syngnathus  fuscus  Storer). 

DIAGNOSIS. — Syngnathids  with  subcaudal 
brood  pouch  protected  by  plates;  nuchal  and 
prenuchal  plates  and  scutella  present;  superior 
ridges  of  trunk  and  tail  discontinuous;  lateral 
ridge  of  trunk  and  superior  ridge  of  tail  contin- 
uous or  subcontinuous;  inferior  trunk  and  tail 
ridges  continuous;  opercular  ridge  present  or  re- 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


199 


-  31                                                    m  4a.  Total  rings  63  or  more,  usually  64-66; 

g  2-    "J____l_i— -— -^ — " p              %"  35-41  dorsal  rays,  usually  37-40  (see  Fig. 

-i  ,.  11),  dark  bar  extending  diagonally  from 

</i     ___^__^_^_^___^_^___^_^__  orbit  across  opercle 

38         «         42         44         46         48         so  ex/7/s  (Osbuni  and  Nichols) 

SLmm  Halfmoon  Bay,  California,  to 

FIGURE  8.     Comparison  of  snout   length   plotted  against  Bahfa  Magdalena,   Baja  Cali- 

standard  length  for  Bryx  veleronis  (closed  circles)  and  Bryx  fornia;  on  sand  bottoms 

darionensis  (closed  squares).  4b.  Total  rings  67-72,  usually  68-70;  39-48 

dorsal  rays,  usually  42-46  (see  Fig.  11), 

no  dark  bar  across  opercle  _. 

duced  to  only  basal  part;  pectora  ,  dorsal,  ana  ,  ,./-      .       .    c. 

califormensis  Storer 

and  caudal  fins  developed;  dorsal  fin  base  not  .,  ....     . 

Bodega  Bay,  California,  to  Ba- 

raised;  first  hypobranchial  reduced  to  absent;  hfa  Santa  Marfa  Baja  Califor. 

remainder  of  branchial  skeleton  present,  without  nia;  associated  with  kelp  beds, 

tooth  plates;  jugular  plates  well  developed,  sep-  often  epipelagic 

arated  by  a  median  plate;  body  ridges  generally  5a.  Pectoral  fins  long,  extending  posteriorly 

smooth  to  finely  serrate,  both  neither  strongly  across  two  full  trunk  rings... 

toothed  nor  serrate;  2-3  infraorbitals;  25-47  dor-  -    macrobrachium  n.sp. 

sal  rays;  2-4  anal  rays;  11-14  pectoral  rays;  10  Tumbes,  Peru,  to  Puerto  Montt, 

caudal  rays;  14-23  trunk  rings;  33-50  tail  rings.  Chlle 

This  genus  contains  some  of  the  largest  species  5b-  Pectoral  fins  short,  never  extending  pos- 

in  the  family;  S.  californiensis  reaches  a  length  tenorlv  across  two  ful1  rm8s  -                 -   6 

of  533  mm  SL.  Marine  and  fresh  water,  cos-  6a-  Total  nn8s  more  than  60 

mopolitan,  but  most  abundant  in  temperate  seas.  6b-  Total  nn8s  60  or  fewer.,.                          .   8 

DISCUSSION  .-Herald  (1943,  1959),  following  7a-  Trunk  length  L5  or  more  in  tai1  length; 

the  preliminary  analysis  of  Duncker  (1915),  di-  total  rm8s  60-68=  dorsal  ravs  33~45 

vided  the  genus  into  four  subgenera  differen-  -  euc"rous  n.sp. 

tiated  on  the  basis  of  brood-pouch  closure.  On  Redondo  Beach,  California,  to 

...                  ,        T              .     ,        ,                       .,  Bahfa  de  Ballenas,  Baja  Cali- 

osteological  grounds,  I  conclude  that  two  of  fornia.  near  bottom  off  rocky 

these  subgenera  (Bryx  and  Microsyngnathus)  shores 

together  constitute  a  distinct  genus  (Bryx).  75.  Trunk  length  1.5  or  less  in  tail  length;  total 

Syngnathus  is  the  most  speciose  pipefish  genus,  rings  53-64;  dorsal  rays  28-43 

with  perhaps  40  species.  leptorhynchus  Girard  (in  part) 

8a.  Interorbital  3  in  postorbital;  head  usually 

Key  to  Eastern  Pacific  Species  of  more  than  g  6  m  SL     leptorhynchus  Girard 

Syngnathus  _    (                             D  , , 

Southeastern  Alaska  to  Bahia 

la.  Trunk  rings  14-16  (x  =   15)    Santa  Maria,  Baja  California; 

.,    aullSCUS  (Swain)  in  eelgrass  of  bays 

Santa  Barbara  Channel,  Cali-  8b-  Interorbital  more  than  3  in  postorbital; 

fornia,  to  northern  Peru;  in  eel-  head  Usually  less  than  8.6  in  SL 

grass  of  bays  and  sloughs  insulae  n.sp. 

lb.  Trunk  rings  17  or  more  (6  of  914  speci-  isia  Guadaiupe,  Mexico,  in  ai- 

mens  of  S.  leptorhynchus  have  16)    2  gae  and  eelgrass,  20-34  m 

2a.   Snout  long,  compressed,  usually  con- 
tained less  than  1.8  in  head  length 3  Syngnathus  auliscus  (Swain) 

2b.  Snout  shorter,  round,  usually  contained  (Figures  4D  &  9) 

more  than  1.8  times  in  head  length...        _    5  Siphostoma  auliscus  SWAIN,   1882:547  (original  description; 

3a.    Total    rings   fewer  than   63;   all    ridges   of  Santa  Barbara  and  San  Diego,  California;  types  lost);  JOR- 

trunk  and  tail  distinctly  keeled DAN  AND  GILBERT  1883:905  (description;  range);  SWAIN 

carinatus  (Gilbert)  AND  MEEK  1884:238  (diagnosed  in  key);  JORDAN  AND  Ev- 
ERMANN   1896:767  (description;  range);  EIGENMANN  AND 

Northern  Gulf  of  California  ElGENMANN  ,892: 144  (San  Diego). 

3b.   Total   rings    more   than   63;   all   ridges   of  Syngnathus  auliscus;  STARRS  AND  MORRIS  1907: 184  (range); 

trunk  and  tail  rounded  __                                 _    4  HUBBS   1916:160  (description);  ULREY  AND  GREELEY 


200 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


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FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


201 


TABLE  5.     NUMBER  OF  SPECIMENS  (N)  AND  STANDARD  LENGTH  RANGE,  TOGETHER  WITH  RANGE  AND  MEAN  (?)  OF  SE- 
LECTED CHARACTERS  EXPRESSED  IN  THOUSANDTHS  OF  STANDARD  LENGTH  IN  Syngnathus. 


Head 

Snout 

Snout 

Body 

Dorsal-base 

Pectoral-fin 

length 

length 

depth 

depth 

length 

length 

Species 

SL 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

Range 

X 

N 

auliscus 

46-134 

98-136 

113 

44-56 

48 

10-19 

15 

18-46 

31 

98-124 

112 

13-21 

18 

9 

carinatus 

1  16-230 

113-141 

128 

65-89 

76 

8-10 

9 

16-26 

22 

114-138 

127 

15-26 

21 

17 

exilis 

75-238 

102-148 

121 

50-88 

68 

7-13 

10 

20-44 

29 

95-120 

107 

13-21 

16 

37 

californiensis 

78-382 

111-143 

120 

55-93 

69 

6-12 

9 

15-41 

24 

100-128 

115 

9-18 

14 

33 

macrobrachium 

92-190 

112-144 

137 

60-82 

72 

10-14 

12 

26-35 

30 

121-140 

130 

21-34 

29 

15 

euchrous  euchrous 

1  1  1-249 

100-132 

114 

47-78 

55 

8-14 

11 

22-40 

29 

111-128 

118 

10-23 

17 

20 

e.  ollotropis 

1  16-244 

93-108 

103 

44-57 

49 

10-12 

11 

22-30 

26 

96-127 

112 

10-15 

13 

10 

leptorhynchus 

57-245 

104-148 

122 

48-87 

65 

8-15 

12 

20-55 

27 

97-141 

118 

10-20 

16 

92 

ins  ula  e 

88-180 

107-117 

111 

46-57 

52 

10-14 

11 

19-30 

24 

105-114 

109 

13-19 

16 

7 

1928:42  (synonymy:  range);  ULREY  1929:6  (Lower  Califor- 
nia); JORDAN,  EVERMANN,  AND  CLARK  1930:240  (range); 
ULREY  1932:77  (Golfo  de  California):  BARNHART  1936:34 
(description;  range);  HERALD  1940:60,  63  (key;  synonymy: 
range);  CASTRO-AGUIRRE  ET  AL.  1970:131  (not  common  in 
Gulf  of  California);  MILLER  AND  LEA  1972:89  (description: 
range):  CHIRICHIGNO-F.  1974:351  (range). 

Singnathus  auliscus;  VAILLANT  1894:70  (Gulf  of  California). 

Siphostoma  sinaloae  JORDAN  AND  STARKS  1896:268  (original 
description;  Mazatlan;  holotype  SU  2945). 

Syngnathus  tweedlei  MEEK  AND  HILDEBRAND,  1923:259  (orig- 
inal description:  Chame  Point,  Panama;  holotype  USNM 
82088). 

DIAGNOSIS. — A  Syngnathus  with  26-33  (27- 
30)  dorsal  rays;  10-14  pectoral  rays;  2-3  anal 
rays;  14-16  (15)  trunk  rings;  34-39  (36-37)  tail 
rings;  0.2-1.5  +  5-7.2  rings  covered  by  dorsal 
fin;  total  rings  covered  by  dorsal  fin  6-7.9;  15- 
21  pouch  rings;  head  7.1-10.1  in  SL;  dorsal  fin 
base  0.8-1.6  in  head;  snout  2.0-2.8  in  head; 
maximum  known  size  178  mm  SL;  smallest  sex- 
ually mature  male  71  mm  SL.  See  Tables  4  and 
5  for  additional  counts  and  measurements. 

DESCRIPTION. — All  ridges  of  body  smooth, 
rounded,  low  but  distinct.  Prenuchal  and  nuchal 
ridges  low  but  distinct.  Opercles  strongly  con- 
vex, with  median  ridge  reaching  half  the  length. 
Snout  ridge  running  full  length  of  snout  to  inter- 
orbital.  Head  not  raised  at  orbit,  so  that  snout 
ridge  almost  horizontal.  Supraorbital  ridge  orig- 
inating just  anterior  to  nostrils  and  ending  above 
anterior  edge  of  opercle.  Pectoral  cover  plate 
with  one  obsolete  ridge.  Lateral  trunk  and  su- 
perior tail  ridges  overlap  for  one  ring.  Slightly 
acute  ventral  trunk  ridge.  Three  lateral  line  pa- 
pillae per  plate.  Dermal  flaps  rarely  present. 

Plates  protecting  brood  pouch  well  developed, 
often  as  deep  as  overlying  tail  rings.  Eggs  0.6- 


0.7  mm  in  greatest  diameter,  arranged  in  two 
layers  four  across.  Brooding  males  collected  in 
April  through  August. 

Fins  of  moderate  size,  caudal  as  long  as  last 
three  tail  rings.  Pectoral  base  about  1.5  in  its 
length. 

Juveniles  less  than  25  mm  SL  (Fig.  9)  are 
much  spinier  than  adults.  Large  broad-based 
spine  on  each  frontal;  opercle  has  a  broad-based 
lateral  projection,  and  each  ring  has  a  well-de- 
veloped spine  on  each  ridge.  Dorsal  fin  high. 
Mouth  large  and  upturned.  Body  whitish  with 
light  brown  pigmented  spots  forming  a  band  dor- 
sal to  gut  anteriorly  and  continuing  posteriorly 
along  ventral  surface  of  tail.  Occasionally  with 
sprinkling  of  pigment  spots  on  body,  concen- 
trated on  ventral  surface  of  gut  and  along  dorsal 
base. 

Coloration  in  life.  Variations  of  brown  or 
green,  never  red. 

Coloration  in  alcohol.  Usually  light  tan  to 
brown,  lighter  ventrally.  Median  caudal  rays 
brown;  outer  two  rays  light  tan;  all  other  fins 
colorless.  Usually  a  dusky  blotch  below  nostrils. 
Sometimes  a  brownish  streak  at  lower-posterior 
corner  of  orbit.  Often  there  is  on  the  body  much 
darker  mottling,  which  may  alternate  with  light 
areas,  especially  posteriorly,  giving  a  dark-light 
banded  appearance.  Lighter  bands  usually  nar- 
rower and  separated  by  five  to  six  rings. 

HABITAT. — Eelgrass  of  bays  and  sloughs.  Oc- 
casionally in  floating  Sargassum. 

COMPARISONS. — Although  S.  auliscus  is  a 
chiefly  tropical  species,  it  occurs  rather  fre- 
quently in  the  southern  part  of  San  Diego  Bay, 
where  the  temperature  is  raised  by  a  thermal 


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B 


FIGURE  9.     Juvenile  Syngnathus  auliscus  from  San  Diego  Bay,  California.  (A)  9.2  mm  SL.  (B)  16.3  mm  SL. 


effluent;  this  is  the  only  place  in  California 
where  this  species  has  been  found  in  larger  num- 
bers than  other  pipefishes.  It  may  be  confused 
in  this  region  with  the  similar  S.  leptorhynchus. 
S.  auliscus  differs  from  5.  leptorhynchus  in  hav- 
ing 14-16  trunk  rings  rather  than  17  or  more;  in 
having  a  deeper  and  more  robust  trunk;  and  in 
reaching  a  much  smaller  size  as  adults. 

RANGE. — Santa  Barbara  Channel,  California, 
to  Paita,  Peru,  and  throughout  the  Gulf  of  Cal- 
ifornia. 

MATERIAL  EXAMINED. — California:  Newport  Bay,  LACM 
W55-257,  1(162).  Penasquitos  Lagoon,  SIO  73-411,  1(178).  San 
Diego,  SIO  H46-94,  2(127-134);  G.  McGowen  personal  col- 
lection, 89(7.4-31.8). 

Mexico:  Bahia  de  Tortugas,  SIO  H48-53,  1(92).  Bahia  Al- 
mejas,  SIO  65-181,  2(98-111).  Cholla  Bay,  CAS  45071,  2(91- 
99).  Puerto  Penasco,  GCRL  V73: 10729,  2(65-82).  Guaymas, 
R.  Behrstock,  personal  collection,  38(29-112).  Sinaloa,  GCRL 
V71:6572,  1(123);  GCRL  V68:2639,  1(81);  GCRL  V71:6565, 
2(46-52).  Mazatlan,  SU  2945,  1(87),  holotype  of  S.  sinaloae. 

Nicaragua:  Corinto,  CAS  6346,  1(74). 

Costa  Rica:  Puntarenas,  GCRL  V69:3526,  3(44-83). 

Panama:  Chame  Point,  USNM  82088,  1(71),  holotype  of  S. 
tweedlei. 

Peru:  Tumbes,  IMARPE  uncat.,  2(7-100).  Paita,  IMARPE 
uncat.,  1(87). 


Syngnathus  carinatus  (Gilbert) 

(Figure  10B) 

Siphostoma  carinatum  GILBERT,  1892:547  (original  descrip- 
tion; 31°31'45"N,  114°19'W;  lectotype  here  designated  as 
SU  240);  JORDAN  AND  EVERMANN  1896:763  (description). 

Syngnathus  carinatum;  BREDER  1928:24  (Isla  Tiburon,  Bahia 
Adair,  San  Felipe). 

Syngnathus  carinatus;  ULREY  1929:6  (Gulf  of  California);  JOR- 
DAN, EVERMANN,  AND  CLARK  1930:240  (range);  ULREY 
1932:77  (Golfo  de  California);  FOWLER  1944:4%  (range). 


Syngnathus  acus;  (not  of  Linnaeus)  HERALD  1940:60  (diag- 
nosed in  key;  range). 

Syngnathus  griseolineatus  leptorhynchus;  (not  of  Girard)  CAS- 
TRO-AGUIRRE  ET  AL.  1970:132  (Gulf  of  California). 

DIAGNOSIS. — A  Syngnathus  with  40-46  (41- 
45)  dorsal  rays;  13-15  pectoral  rays;  3-4  anal 
rays;  16-18  (17)  trunk  rings;  39-43  (41-42)  tail 
rings;  56-60  total  rings;  dorsal  on  0.1-1. 8  +  7.5- 
11.1  rings;  total  rings  covered  by  dorsal  9- 
12.1;  21.5-22  rings  covered  by  brood  pouch; 
head  7.1-8.6  in  SL;  dorsal  fin  base  0.9-1.2  in 
head;  snout  1.6-1.8  in  head;  maximum  known 
size  230  mm  SL;  smallest  sexually  mature  male 
182  mm  SL.  See  Tables  4  and  5  for  additional 
counts  and  measurements. 

DESCRIPTION. — All  ridges  on  body  usually 
well  developed  and  serrate.  Supraoccipital,  nu- 
chal, and  prenuchal  ridges  low  and  finely  ser- 
rate. Opercular  and  supraorbital  ridges  low. 
Snout  ridge  slightly  elevated  and  serrate,  ex- 
tending posteriorly  to  region  above  anterior  edge 
of  orbit.  Pectoral  cover  plate  with  only  superior 
ridge.  Superior  trunk  ridge  extending  forward 
dorsally  to  gill  opening  and  ending  just  below 
nuchal  plate.  Lateral  trunk  and  superior  tail 
ridges  usually  not  overlapping,  but  may  overlap 
for  half  width  of  last  trunk  ring.  Scutella  easily 
discernible,  about  half  as  deep  as  median  trunk 
plate.  Lateral  line  consisting  of  6-7  papillae  per 
plate.  No  dermal  flaps. 

Protecting  plates  of  brood  pouch  well  devel- 
oped, more  than  half  as  high  as  adjacent  tail;  mar- 
gin of  folds  of  brood  pouch  smooth.  Eggs  0.9- 
1.0  mm  in  greatest  diameter,  arranged  in  two 
layers  six  across.  Males  with  eggs  have  been 
collected  in  November. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


203 


B 


FIGURE  10.  Anterior  part  of  body  of  four  species  of  eastern  Pacific  Syngnathidae.  (A)  Syngnathus  macrobrachium,  holotype, 
USNM  176501.  (B)  Syngnathus  carinatus,  a  200-mm-SL  female,  SIO  72-60.  (C)  Syngnathus  exili.s,  a  238-mm-SL  female,  SIO 
75-661.  (D)  Syngnathus  californiensis,  a  285-mm-SL  female,  SIO  62-552. 


Fins  all  well  developed.  Pectoral  fin  extends 
posteriorly  to  edge  of  first  trunk  ring  or  middle 
of  second  trunk  ring. 

Coloration  in  alcohol.  Older  specimens  a  uni- 
form light  tan  to  dark  brown,  with  caudal  dusky 
and  dark  spots  at  bases  of  dorsal  rays.  Fresh 
specimens  with  immaculate  white  undersides 
becoming  silvery  on  opercle  and  pectoral  cover 
plate.  Dorsum  light  green-brown  with  white 
areas  around  scutella,  which  may  be  connected 
by  white  streaks  to  adjacent  scutella.  Thus,  a 
chainlike  pattern  often  evident  dorsolaterally. 
Caudal  fin  green-brown  centrally  with  light 


edges.  Snout  with  the  same  green-brown  color 
dorsally.  Other  fins  colorless. 

MERISTIC  VARIATION. — The  number  of  trunk 
rings  in  5.  carinatus  is  relatively  stable,  usually 
17,  except  that  three  of  the  four  specimens  from 
Bahfa  Kino  had  16.  The  same  trend  is  shown  by 
the  number  of  tail  rings.  The  normal  range  is 
from  41-42,  but  two  of  the  four  specimens  from 
Bahfa  Kino  had  40  tail  rings. 

COMPARISONS. —  As  indicated  in  the  account 
of  S.  macrobrachium,  S.  carinatus  is  closely 
related  to  macrobrachium,  schlegeli,  andfollet- 
ti.  It  seems  to  be  most  closely  related  to  the 


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Japanese  5.  schlegeli.  It  differs  from  schlegeli 
in  coloration  (light  coloration  with  chainlike  pat- 
tern rather  than  dark  brown),  longer  snout,  and 
in  that  the  superior  trunk  ridge  is  continued  an- 
teriorly above  the  gill  opening.  The  relationship 
of  S.  carinatus  to  a  Japanese  fish  parallels  the 
distribution  indicated  by  Dawson  (1944)  for 
some  algae. 

RANGE. — Northern  part  of  Gulf  of  California, 
from  head  of  Gulf  and  from  Bahfa  San  Luis  Gon- 
zaga  in  the  west  to  Bahfa  Kino  in  the  east. 

MATERIAL  EXAMINED. — Gulf  of  California:  Mouth  of  Rio 
Colorado,  SIO  63-484,  2(132-139).  Between  El  Golfo  and  San 
Felipe,  S.  Guevara  personal  collection,  20(161-216).  Off  Bahfa 
Adair,  BOC  1184,  11(86-148).  31°32'30"N,  114°20'W,  USNM 
48250,  3(185-287);  USNM  127139,  2(187-189);  USNM  125028, 
2(178-183),  paralectotypes.  31°31'45"N,  114°19'W,  SU  240, 
1(198),  lectotype;  SU  67670,  3(186-201),  paralectotypes.  San 
Felipe,  SIO  72-60,  2(203-209);  SIO  58-164,  1(127):  SIO  H47- 
53,  47(132-165);  SIO  74-118,  3(182-230);  BOC  1017,  5(81- 
156).  Punta  Diggs,  UCLA  W62-61,  5(131-152).  Isla  Willard, 
LACM  22298,  1(210).  Isla  Tiburon,  BOC  1010,  1(123).  Bahfa 
Kino,  SIO  60-1 13,  4(1 15-133). 

Syngnathus  exilis  (Osburn  and  Nichols) 

(Figures  IOC  &  11) 

Siphostoma  exile  OSBURN  AND  NICHOLS,  1916:153  (original 

description;  west  San  Benito  Island,  Baja  California:  holo- 

type  USNM  87543,  formerly  AMNH  5203). 
Syrictes  exilis;  JORDAN,  EVERMANN,  AND  CLARK  1930:243 

(range). 
Syngnathus  californiensis  californiensis',  HERALD  1940:60  (in 

part);  1941:62  (in  part). 
Syngnathus  exilis;  ULREY  1929:6  (listed);  MILLER  AND  LEA 

1972:212  (range). 

DIAGNOSIS. — A  Syngnathus  with  36-41  (rare- 
ly 35,  42  or  43)  (37-40)  dorsal  rays;  11-14  pec- 
toral rays;  3  anal  rays;  17-21  (18)  trunk  rings; 
43-49  (rarely  50)  (45-47)  tail  rings;  63-68  (64- 
66)  total  rings;  dorsal  on  0-1.9  +  7-9.5  rings; 
brood  pouch  covering  19-26  rings;  head  6.8-9.8 
in  SL;  dorsal  fin  base  0.9-1.4  in  head;  snout  1.6- 
2.0  in  head;  eggs  1.0-1.2  mm  in  greatest  diam- 
eter, arranged  in  two  layers  four  across.  Maxi- 
mum known  size  253  mm  SL;  smallest  sexually 
mature  male  125  mm  SL.  See  Tables  4  and  5  for 
additional  counts  and  measurements. 

DESCRIPTION. — Ridges  of  trunk  and  tail 
smooth  and  rounded.  Nuchal  and  prenuchal 
ridges  obsolete  to  absent,  smooth.  Opercular 
ridge  distinct,  reduced  to  basal  third  of  opercle. 
Snout  ridge  low,  smooth  to  slightly  serrate, 
reaching  from  vomer  to  anterior  part  of  inter- 
orbital;  slightly  elevated  posteriorly.  Superior 
ridge  of  pectoral  cover  plate  absent,  inferior 


ridge  obsolete.  Lateral  trunk  and  superior  tail 
ridges  overlap  on  first  tail  ring.  Trunk  and  tail 
ridges  smooth  to  slightly  rugose.  Scutella  small, 
about  one-third  as  deep  as  adjoining  plates. 
Snout  moderately  compressed.  Abdomen  typi- 
cally and  rather  distinctly  with  posterior  swell- 
ing, especially  in  juveniles  and  females.  Lateral 
line  consisting  of  4-5  papillae  per  plate.  Often 
with  a  pair  of  stitches  on  first  trunk  ring  and 
nuchal  plates.  No  dermal  flaps. 

Pouch  protecting  plates  well  developed,  about 
as  high  as  adjacent  tail.  Eggs  1.0-1.2  mm  in 
greatest  diameter,  arranged  in  two  layers  four 
across.  Brooding  males  found  throughout  year. 

Pectoral  fins  reach  posteriorly  to  just  past 
junction  of  first  and  second  trunk  rings.  Caudal 
fin  as  long  as  last  three  tail  rings. 

Coloration  in  life.  From  C.  L.  Hubbs's  field 
notes  for  SIO  H47-43. 

The  specimens  become  sooty  toward  the  caudal.  Of  the  half 
grown,  one  is  blackish  green  the  other  tan,  much  variegated 
with  lighter  and  darker.  All  are  much  speckled  with  light  on 
top  of  head.  The  female  varies  from  greenish  brown  ante- 
riorly to  reddish  brown  posteriorly.  The  male  is  brown,  be- 
coming orange  toward  the  brood  pouch.  The  lower  part  of 
the  opercle  of  all  is  silvery  with  a  more  or  less  distinct  dark 
stripe  behind  the  lower  part  of  the  eye.  Dark  markings  on 
lower  part  of  opercle  are  incipient.  All  but  the  adult  male 
have  a  diffuse  dark  streak  on  belly.  The  adults  show  the 
half  crescents  on  the  body  just  above  the  ventrolateral  keel. 
The  adult  male  is  rather  conspicuously  banded  with  light. 
The  others  indefinitely  so. 

Coloration  in  alcohol.  The  most  striking  fea- 
ture of  alcoholic  specimens  is  the  dusky  stripe 
on  the  snout  extending  from  the  nostrils  ante- 
riorly on  the  greater  part  of  the  snout,  and  the 
bar  that  courses  diagonally  from  the  postero- 
ventral  margin  of  the  orbit  to  the  opercle.  These 
markings  may  be  wanting  in  older  specimens. 
The  general  body  color  is  typical  for  most 
species  of  Syngnathus;  i.e.,  a  ground  color  tan 
to  dark  brown,  with  various  mottlings  of  darker 
brown. 

HABITAT. — On  the  bottom  off  sandy  beaches, 
often  associated  with  pieces  of  detached  algae. 
Occasionally,  juveniles  are  taken  in  the  plank- 
ton, sometimes  associated  with  flotsam.  Two 
specimens  (SIO  H5 1-203  and  CAS  32983)  were 
tiaken  from  the  guts  of  albacore  (Thunnus  alal- 
unga). 

COMPARISONS. — S.  exilis  is  a  long-snouted 
pipefish  of  the  outer  coast,  resembling  5.  cali- 
forniensis in  general  appearance.  It  can  be  dis- 
tinguished from  that  species  by  its  smaller  size 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


205 


at  maturity,  generally  fewer  trunk  rings  (Jc  = 
18.3  vs  x  =  20.8),  coloration  (especially  the 
presence  of  the  dark  oblique  bar  at  the  posterior 
corner  of  the  orbit),  the  abdomen  usually  with 
a  posterior  swelling,  and  the  sand-bottom  vs 
kelp-bed  habitat.  Figure  1 1  provides  information 
for  easy  separation  of  S.  exilis  from  S.  califor- 
niensis. 

RANGE. — Half  Moon  Bay,  California,  to  Ba- 
hia  Magdalena,  Baja  California,  including  the 
channel  islands  of  California  and  Isla  Guada- 
lupe,  Mexico. 

MATERIAL  EXAMINED. — California:  Half  Moon  Bay,  SIO 
54-106,  1(217).  Guide  Seamount,  CAS  32983,  1(97).  Del  Monte 
Beach,  SIO  74-195,  2(92-1%).  Monterey,  CAS  27950,  1(89). 
San  Luis  Obispo,  SIO  55-35,  1(164);  SIO  H50-195,  2(217-227). 
Los  Angeles,  SCCWRP  sta.  T,-75,  4(176-199);  SCCWRP  sta. 
T-0,  4(132-203).  Long  Beach,  UCLA  W50-128,  2(228-235). 
Newport  Beach,  SIO  H45-70,  17(120-239).  Dana  Point, 
SCCWRP  sta.  Fv.  3,  5(105-190).  Torrey  Pines  Lagoon, 
UCLA  W63-49,  2(170-175).  La  Jolla,  SIO  H47-43,  4(96-163); 
SIO  H47-106,  48(107-242);  SIO  H51-153,  13(136-227);  SIO 
H46-63,  large  series;  SIO  H47-158,  34(96-225);  SIO  H47-134, 
1(170);  SIO  H48-163,  large  series.  Santa  Cruz  Island,  SIO 
H51-259,  1(1 15).  Santa  Catalina  Island,  LACM  30510-9,  1(85); 
LACM  30865-3,  1(42).  Mission  Bay,  SIO  61-412,  1(113);  San 
Diego  Bay,  SIO  H50-112,  1(219). 

Baja  California:  Islas  Coronados,  SIO  H46-125,  1(145).  Pun- 
ta  Banda  and  Isla  de  Todos  Santos,  SIO  61-284,  1(120);  SIO 
68-302,  1(160);  SIO  H51-24,  2(102-118).  Punta  Santo  Tomas, 
SIO  65-440,  1(103).  Cabo  Colnett,  SIO  H51-303,  1(181);  CAS 
3295,  8(114-161).  Bahia  San  Quintin,  SIO  H50-25,  1(239);  SIO 
60-453,  1(135).  Bahi'a  San  Carlos,  SIO  H52-210,  4(114-247). 
Bahia  Blanca,  SIO  60-327,  3(158-250).  Bahi'a  Playa  Maria, 
SIO  H52-169,  2(120-224).  Isla  Guadalupe,  SIO  65-72,  1(130). 
Bahi'a  Santa  Rosalia,  SIO  62-282,  1(164).  Bahi'a  Sebastian  Viz- 
cafno,  SIO  H52-135,  9(131-228).  Islas  San  Benitos,  USNM 
87534,  1(166),  holotype.  Isla  Cedros,  SIO  68-345,  1(175); 
UCLA  W60-126,  1(123);  SU  67915,  1(117);  SIO  65-591,  1(220); 
SIO  62-756,  1(145);  SIO  H53-110,  1(193).  Punta  Eugenia,  SIO 
61-398,  1(70).  Cabo  Tortolo,  SIO  H48-55,  1(180).  Bahi'a  Tor- 
tugas,  SIO  H48-51,  1(98);  SIO  63-806,  1(141);  SIO  59-25, 
1(155):  SIO  H48-33,  3(192-238);  SU  2488,  1(123);  AMNH 
5588,  5(99-118),  paratypes.  Bahi'a  San  Roque,  SIO  63-16, 
1(84);  SIO  64-185,  1(152).  Bahia  Asuncion,  SIO  68-303,  1(161); 
SIO  62-535,  2(100-110):  SIO  68-301,  2(96-151);  SIO  H51-96, 
1(174);  SIO  H48-92,  1(115);  SIO  66-147,  1(190);  SIO  64-68, 
3(161-226).  Bahi'a  San  Hippolito,  SIO  64-750,  14(70-253).  Ba- 
hia de  Ballenas,  SIO  64-405,  1(173);  SIO  64-765,  1(180);  SU 
767,  1(147).  Bahi'a  Santa  Maria,  SIO  H50-149,  1(136).  Bahi'a 
Magdalena,  SIO  H50-96,  1(67). 

Syngnathus  californiensis  Storer 

(Figures  10D,  11,  16C) 

Syngnathus  californiensis  STORER,  1848:73  (original  descrip- 
tion; California;  holotype  lost);  STARKS  AND  MORRIS 
1907:183  (meristic  features);  STARKS  1911:177  (not  included 
in  Puget  Sound  fauna);  HUBBS  1916:158  (description); 
1921:4  (range;  counts);  ULREY  AND  GREELEY  1928:42  (syn- 
onymy: range);  ULREY  1929:6  (included  in  list);  JORDAN, 


EVERMANN,  AND  CLARK  1930:240  (range);  BARNHART 
1936:34  (description);  HERALD  1940:64  (in  part;  diagnosed 
in  key);  1941:62  (description;  range:  synonymy);  MILLER 
AND  LEA  1972:89  (description;  range). 

Siphostoma  californiense;  JORDAN  AND  GILBERT  1881:453 
(synonymy);  JORDAN  AND  JOUY  1881:1  (Santa  Barbara, 
Monterey,  and  San  Francisco);  JORDAN  AND  GILBERT 
1882:69  (in  part;  5.  griseolineatus  included);  SWAIN 
1882:308  (description;  synonymy);  JORDAN  AND  GILBERT 
1883:384  (description;  synonymy;  range);  SWAIN  AND 
MEEK  1884:238  (diagnosed  in  key):  JORDAN  AND  EVER- 
MANN 1896:764  (description;  synonymy;  range). 

Siphostoma  californiensis;  EVERMANN  AND  LATIMER  1910:136 
(San  Pablo  Bay,  California,  probably  5.  leptorhynchus). 

DIAGNOSIS. — A  Syngnathus  with  40-48  (42- 
46)  dorsal  rays;  12-14  pectoral  rays;  3  anal  rays; 
19-22  (21)  trunk  rings;  46-52  (48-49)  tail  rings; 
66-71  total  rings;  dorsal  on  0.2-1.9  +  7.5-9.5 
rings;  total  rings  covered  by  dorsal  8.5-10.5; 
19.5-25  rings  covered  by  brood  pouch;  head 
6.2-8.9  in  SL;  dorsal  fin  base  0.9-1.2  in  head; 
snout  1.5-2.0  in  head;  maximum  known  size  533 
mm  SL;  smallest  mature  male  191  mm  SL.  See 
Tables  4  and  5  for  additional  counts  and  mea- 
surements. 

DESCRIPTION. — All  ridges  on  body  distinct, 
smooth,  and  rounded.  Supraoccipital,  nuchal, 
and  prenuchal  ridges  absent.  Opercular  ridge  re- 
duced to  absent.  Snout  ridge  low,  not  elevated, 
and  smooth,  extending  posteriorly  to  interorbit. 
Supraorbital  ridge  absent.  Pectoral  cover-plate 
ridges  absent.  Superior  trunk  ridge  indistinct  an- 
terior to  pectoral  fin.  Lateral  trunk  ridge  and 
superior  tail  ridge  overlap  on  first  tail  ring.  Scu- 
tella  small,  about  one-third  depth  of  adjoining 
plates.  Lateral  line  consisting  of  7-8  papillae  per 
plate.  No  dermal  flaps. 

Protecting  plates  of  brood  pouch  well  devel- 
oped, about  as  high  as  half  adjacent  tail;  margin 
of  brood  pouch  folds  smooth.  Eggs  1.1-1.3  in 
greatest  diameter,  arranged  in  2  layers  3-4 
across.  Males  with  young  or  eggs  have  been  col- 
lected in  July-October. 

The  fins  are  normally  developed.  Pectoral  fin 
reaching  posteriorly  just  past  junction  of  first 
and  second  trunk  rings.  Caudal  fin  as  long  as  last 
three  tail  rings. 

Coloration  in  life.  From  C.  L.  Hubbs's  field 
notes  for  SIO  H45-69. 

Belly  generally  yellow-green,  sometimes  pale  blue  or  even 
pale  brownish.  Especially  conspicuous  are  the  small  dark 
speckles,  occasionally  weak  or  absent,  along  the  carinae  of 
the  trunk  between  the  plates.  These  provide  an  almost  in- 
variable distinction.  The  streak  along  the  mid-line  of  the 
belly  is  lacking  or  very  weakly  developed.  In  almost  all 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


46- 

44- 

42- 

W40- 


B 


63      65        67 
TOTAL  RINGS 


71 


<J 

o       8 

8 

B        °   \  • 

<r 

-I38' 

8        D     \ 

B 

8 

§        B 

w 

oe36- 

a 

8 

o 

0 

a 

, 

C 

64  66 

TOTAL    RINGS 


68 


70 


72 


FIGURE  11.  Comparison  of  meristic  characters  between  Syngnathus  exilis  and  Syngnathus  calif orniensis .  (A)  Frequency 
of  dorsal  ray  counts.  5.  exilis  vertical  lines,  5.  californiensis  diagonal  lines.  (B)  Frequency  of  total  ring  number.  S.  exilis 
vertical  lines,  S.  californiensis  diagonal  lines.  (C)  Number  of  dorsal  rays  plotted  against  total  number  of  rings.  5.  exilis  open 
squares,  5.  californiensis  dots. 


specimens  the  light  streakings  in  the  upper  and  lateral  as- 
pects of  the  head  and  body  form  more  or  less  definite  and 
extensive  wavy  lengthwise  lines,  typically  forming  a  ring  on 
the  first  nuchal  plate.  When  light  markings  are  developed 
along  the  ventral  carina  of  trunk  (in  about  one-third  of  spec- 
imens) there  tend  to  be  more  definite  elliptical  or  oval  marks 
more  or  less  separated  from  the  light  color  of  the  belly,  and 
when  they  are  developed  similar  marks  are  usually  evident 
along  the  median  and  dorsal  carinae. 

Coloration  in  alcohol.  The  young,  which  are 
associated  with  drifting  kelp,  often  are  light  gray 
to  tan  with  a  dark  streak  alongside  the  snout. 

Adults  vary  from  tan  to  brown,  often  with  in- 
distinct darker  blotches  and  wavy  lines  about 
the  head  and  opercle;  especially  pronounced 
along  the  sides  of  the  trunk.  The  dark  snout 
stripe  most  often  remains  as  a  dark  area  around 


and  below  the  nostrils.  The  caudal  has  the  same 
color  as  the  body.  The  remaining  fins  are  col- 
orless except  for  melanophores  along  the  rays. 

HABITAT. — Offshore  kelp  (Macrocystis)  beds, 
and  frequently  associated  with  drifting  kelp. 

COMPARISONS. — S.  californiensis  is  a  very 
distinctive  fish  as  an  adult.  The  combination  of 
large  size  and  high  number  of  rings  and  dorsal 
rays  serves  to  distinguish  5.  californiensis  from 
all  other  California  pipefishes.  S.  euchrous  most 
closely  approaches  5.  californiensis  in  size  and 
number  of  rings,  but  its  short  round  snout  easily 
distinguishes  it  from  the  longer  compressed 
snout  of  S.  californiensis.  S.  californiensis  ju- 
veniles are  easily  confused  with  adults  of  S.  ex- 
ilis (see  comparison  in  account  of  5.  exilis). 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


207 


RANGE. — Bodega  Bay,  California,  south  to 
Bahfa  Santa  Maria,  Baja  California,  and  includ- 
ing the  offshore  islands  (essentially  the  same 
distribution  as  that  of  the  giant  kelps  Macrocys- 
tis  pyrifera  and  M.  angustifolia). 

MATERIAL  EXAMINED. — California:  Bodega  Bay,  CAS 
32933,  1(162).  San  Francisco,  CAS  60233,  2(154-171);  USNM 
27199,  1(401).  Moss  Landing,  SIO  62-510,  1(284);  SIO  53-194, 
3(131-192).  Monterey  Bay,  SU  19261,  1(128);  SU  36465,  6(94- 
325);  SU  5327,  3(157-184);  CAS  13409,  1(177);  SU  58461, 
3(57-84).  Guide  Sea  Mount,  CAS  32983,  1(98).  36°N,  123°W, 
LACM  30458-1,  1(212).  W.  R.  Hearst  State  Park,  SIO  73-301, 
13(174-254).  Morro  Bay,  UCLA  W50-196,  2(162-185).  Diablo 
Cove,  LACM  31300-24,  2(160-203).  Avila  Bay,  SIO  H50-195. 
Santa  Barbara,  SIO  61-426,  30(119-362);  SIO  H44-12,  1(173); 
SIO  H5 1-128,  3(119-327).  Santa  Rosa  Island,  UCLA  W50-99, 
2(161-171).  Santa  Cruz  Island,  UCLA  W50-87,  1(247);  UCLA 
W50-97,  1(183).  San  Pedro,  SIO  H44-38.  Long  Beach,  CAS 
31512,  1(241).  Del  Mar,  SIO  58-107,  2(400-410).  Catalina  Ba- 
sin, S.  Imsand  personal  collection,  16(78-100).  La  Jolla,  SIO 
H52-181,  1(115);  SIO  H50-7,  1(437);  SIO  H47-153,  1(172).  Bird 
Rock,  SIO  75-427,  1(533).  Mission  Beach,  SIO  67-29,  1(530). 
Mission  Bay,  SIO  H45-8,  5(121-184);  CAS  11428,  1(122);  SIO 
H44-28,  1(125);  SIO  H46-75,  2(168-180).  Point  Loma,  SIO 
H51-347,  7(151-387). 

Mexico:  Bahfa  de  Todos  Santos,  SIO  H53-229,  2(223-269); 
SIO  61-284,  13(111-323);  SIO  64-806,  5(125-349).  Punta  Ban- 
da,  SIO  59-302,  1(194).  Cabo  Colnett,  SIO  H50-62,  1(222).  Isla 
San  Martin,  SIO  H52-219,  1(89);  SIO  H52-218,  3(104-162). 
Bahfa  San  Quintin,  SIO  H51-108,  1(141).  Punta  Baja,  SIO 
H51-401,  1(198).  Isla  San  Geronimo,  SIO  H52-159,  2(190- 
252).  Bahfa  San  Carlos,  SIO  H52-210,  16(175-282).  Punta  Eu- 
genia, SIO  H52-118,  1(125).  Isla  Cedros,  CAS  32952,  1(224). 
Bahfa  Thurloe,  CAS  15424,  1.  Bahfa  Santa  Marfa,  LACM 
W5 1-264,  1(234). 

Syngnathus  macrobrachium  n.sp. 

(Figure  104) 

Syngnathus  acicularis  (not  of  Jenyns);  DELFIN  1901:42 
(range);  HILDEBRAND  1946:155  (in  part;  description;  syn- 
onymy; range);  MANN  1954:190  (description;  range): 
KOEPCKE  1962:200  (range):  DEBUEN  1963:91  (synonymy: 
description);  MORROW  1957:228  (Peru);  CLEMENS  AND 
NOWELL  1963:262  (night  light  off  Peru);  CHIRICHIGNO-F. 
1974:189,  351  (diagnosed  in  key;  range). 

Siphostoma  aciculare  (not  of  Jenyns);  EVERMANN  AND  RAD- 
CLIFFE  1917:53  (synonymy:  description);  NICHOLS  AND 
MURPHY  1922:506  (Chincha  Island). 

Syngnathus  acus  (in  part,  not  of  Linnaeus);  HERALD  1940:60 
(diagnosed  in  key):  FOWLER  1944:496  (range). 

DIAGNOSIS. — A  Syngnathus  with  36-44  (39- 
42)  dorsal  rays;  12-15  pectoral  rays;  2-4  anal 
rays;  16-18  (17-18)  trunk  rings;  40-43  (41-42) 
tail  rings;  57-62  total  rings;  dorsal  on  0.5-2.3 
+  7-9  rings;  total  rings  covered  by  dorsal  fin  9- 
10.3;  17-21  pouch  rings;  head  6.6-8.2  in  SL; 
dorsal  fin  base  0.9-1.2  in  head;  snout  1.8-2.0  in 
head;  maximum  known  size  255  mm  SL;  small- 


est mature  male  84  mm  SL.  See  Tables  4  and  5 
for  additional  counts  and  measurements. 

DESCRIPTION. — All  ridges  of  body  rounded 
and  smooth.  Nuchal  and  prenuchal  ridges  low, 
smooth.  Opercular  ridge  extending  posteriorly 
about  one-fourth  length  of  opercle.  Supraor- 
bital  and  snout  ridges  low,  smooth.  Ridges  of 
pectoral  cover  plate  obsolete.  Trunk  and  tail 
ridges  smooth,  without  indentations  between 
rings.  Superior  trunk  ridge  does  not  extend  an- 
teriorly past  gill  opening.  Scutella  moderately 
developed,  about  equal  in  depth  to  adjacent 
plate.  Lateral  line  consisting  of  4-5  papillae  per 
plate.  No  dermal  flaps. 

Protecting  plates  of  brood  pouch  about  as  high 
as  adjacent  tail;  margin  of  brood  pouch  folds 
smooth.  Egg  size  unknown.  Males  with  embryos 
and  with  recently  emptied  pouches  have  been 
taken  from  October  through  April. 

The  fins  are  well  developed.  The  pectoral  is 
particularly  large,  extending  posteriorly  over 
two  full  rings. 

Coloration  in  alcohol.  Generally  a  light  brown 
to  dusky  with  caudal  fin  darker.  A  dark  streak 
on  mid-ventral  snout,  another  just  anterior  to 
orbit.  Dorsal  and  pectoral  fins  colorless,  with 
chromatophores  present  along  sides  of  individ- 
ual rays. 

MERISTIC  VARIATION. — There  is  little  varia- 
tion in  the  number  of  trunk  rings.  There  is  some 
indication  that  the  Chilean  population  is  char- 
acterized by  having  18  trunk  rings  and  the  Pe- 
ruvian by  17. 

There  is  a  greater  variation  in  the  number  of 
tail  rings.  The  range  for  Peruvian  fish  is  usually 
40-42,  whereas  the  counts  of  the  Chilean  fish 
are  fairly  stable  at  42. 

The  increase  in  number  of  rings  with  higher 
latitudes  is  not  found  in  the  count  of  dorsal  rays. 
The  Peruvian  fish  have  40-42  dorsal  rays, 
whereas  those  from  Chile  have  39-40,  if  the  fish 
from  Hornitos  and  Valparaiso,  Chile,  are  exlud- 
ed. 

COMPARISONS. — S.  macrobrachium  is  very 
close  to  5.  carinatus  and  S.folletti  of  the  west- 
ern Atlantic.  It  differs  from  S.  carinatus  in  hav- 
ing a  longer  snout  (1.6-1.8  rather  than  1.8-2.0 
in  head),  coloration  dark  rather  than  lighter 
gray,  and  a  longer  pectoral  fin  (extending  pos- 
teriorly over  two  full  rings  rather  than  one  to 
one  and  one-half)-  It  differs  from  S.  folletti  in 
having  longer  pectoral  fins,  36-44  dorsal  rays 


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rather  than  37-38,  and  in  having  a  longer  head 
(6.6-8.2  in  SL  rather  than  8.2-9.9). 

RANGE. — Tumbes,  Peru,  south  to  Puerto 
Montt,  Chile. 

ETYMOLOGY. — From  the  Greek  makros — 
long,  and  brachion — arm,  in  reference  to  the 
long  pectoral  fin. 

MATERIAL  EXAMINED.— Holotype:  USNM  176501,  1(225), 
a  mature  male  collected  at  Huasco,  Chile  (28°28'S,  71°14'W), 
on  9-10  Oct.  1844,  by  M.  J.  Lobell. 

Paratypes:  UCLA  W59-49,  1(153),  night  light  off  Punta  Be- 
rejo,  Peru  (10°34'S,  77°56'W),  16  Sep.  1959,  by  J.  Seapin.  SIO 
72-168,  2(177-190),  off  Hornitos,  Chile,  5  m,  otter  trawl,  27 
Apr.  1972,  by  local  fishermen  aboard  the  Luz  III.  USNM 
213485,  4(193-255),  collected  with  holotype.  UCLA  W58-116, 
3(92-103),  bait  net  off  Ilo,  Peru  (17°38'S,  71°18'W),  15  Dec. 
1957,  by  J.  Seapin.  IMARPE  uncat.,  2(122-186),  Zona  de 
Lima,  Peru,  July  1973,  by  K.  Freyman.  IMARPE  uncat., 
10(123-161),  Cantolao  la  Punta,  Peru,  27  Apr.  1964,  by  B.  M. 
Vildoso.  EBMCh  10108-10109,  2(96-111),  Iquique,  Chile, 
Dec.  1957. 

ADDITIONAL  MATERIAL:  Peru:  Cabo  Blanco,  UCLA  W59- 
46,  3(52-60).  Callao,  UCLA  W59-48,  7(74-89).  Lima,  CAS 
45073,  4(116-176).  Bahia  Chilca,  USNM  127856,  3(89-175); 
USNM  127855,  1(107). 

Chile:  Valparaiso,  CAS  12017,  1(126). 

Syngnathus  euchrous  n.sp. 

Syngnathus  californiensis  californiensis  (in  part,  not  of  Sto- 
rer);  HERALD  1940:64;  1941:62. 

DIAGNOSIS. — A  Syngnathus  with  33-45  dor- 
sal rays;  11-14  pectoral  rays;  3  anal  rays;  18-20 
trunk  rings;  41-49  tail  rings;  60-68  total  rings; 
dorsal  on  0.2-1.5  +  6-9  rings;  brood  pouch  cov- 
ering 20-23  rings;  head  8.3-11.2  in  SL;  dorsal 
fin  base  0.8-1.3  in  head;  snout  1.8-2.5  in  head. 

DESCRIPTION. — All  ridges  of  body  smooth  to 
slightly  rugose.  Nuchal  and  prenuchal  ridges  low 
to  obsolete.  Opercle  strongly  convex,  with  weak 
ridge  restricted  to  anterior  fourth  of  opercle. 
Snout  ridge  development  different  in  the  two 
subspecies.  Supraorbital  ridge  reduced.  Inter- 
orbital  flat  to  slightly  concave.  Pectoral  cover 
plate  with  superior  ridge  obsolete  to  absent.  Lat- 
eral trunk  ridge  ending  on  last  trunk  or  first  tail 
ring.  Lateral  line  papillae  5-8  per  plate.  No  der- 
mal flaps. 

Protecting  plates  of  brood  pouch  varying  from 
about  one-third  as  high  to  about  as  high  as  ad- 
jacent tail.  Eggs  1.2  mm  in  greatest  diameter, 
arranged  in  2-4  layers  4-6  across.  Males  with 
eggs  and  young  collected  from  February  through 
August. 

COMPARISONS. — S.  euchrous  is  most  closely 
related  to  S.  leptorhynchus  from  which  it  can  be 
distinguished  by  the  characters  given  in  the  key. 


ETYMOLOGY. — From  the  Greek  eu — well, 
and  chroa — the  color  of  the  skin,  in  reference 
to  the  well-marked  coloration. 

Key  to  the  Subspecies  of 
Syngnathus  euchrous 

la.  Dorsal  rays  usually  39-42;  tail  rings  usu- 
ally 45-46;  21-23  rings  covered  by  brood 
pouch euchrous 

Redondo  Beach,  California  to 
Punta  Eugenia,  Baja  California 

lb.  Dorsal  rays  usually  36-38;  tail  rings  usu- 
ally 42-44;  20-21  rings  covered  by  brood 
pouch.  ollotropis  n.subsp. 

Bahia  Tortugas  to  Bahia  de 
Ballenas,  Baja  California 

Syngnathus  euchrous  euchrous 

(Figure  \2A) 

DIAGNOSIS. — An  S.  euchrous  with  38-45  (39- 
42)  dorsal  rays;  11-14  pectoral  rays;  3  anal  rays; 
18-20  (19)  trunk  rings;  42-49  (45-46)  tail  rings; 
61-68  total  rings;  snout  ridge  concave  in  outline, 
smooth;  maximum  known  size  296  mm  SL; 
smallest  sexually  mature  male  128  mm  SL.  See 
Tables  4,  5,  and  6  for  additional  counts  and  mea- 
surements. 

Coloration  in  life.  From  C.  L.  Hubbs's  field 
notes  for  SIO  H47-43: 

Four  are  rich  brown,  grading  from  greenish  brown  to  red- 
dish brown  with  very  conspicuous  pearly  cross-blotching. 
There  are  a  variable  number  of  blue-white  specks.  The 
crown  and  nape  regions  are  more  or  less  completely  cov- 
ered with  fine  to  coarse  speckles.  Along  the  ventrolateral 
keel  there  are  single  to  compound  specks  on  the  plates 
....  The  opercles  are  flecked  with  silver-blue  and  a  more 
or  less  distinct  wedge  of  the  same  color  extends  downward 
and  backward  from  eye.  The  dorsal  fin  is  entirely  or  almost 
entirely  immaculate. 

Coloration  in  alcohol.  As  is  true  with  most 
species  of  Syngnathus,  excessive  variability 
precludes  using  color  patterns  as  diagnostic  fea- 
tures. A  dusky  patch  or  streak  below  the  nares. 
A  series  of  small  whitish  spots  over  the  entire 
dorsal  surface  of  head  and  body,  often  extending 
onto  sides.  Protecting  plates  of  brood  pouch 
chocolate-brown.  Caudal  brownish  with  white 
on  tips  of  outermost  rays.  Dorsal  and  pectoral 
fins  translucent,  with  some  pigmentation  along 
rays. 

HABITAT. — Near  bottom  off  rocky  shores. 

RANGE. — Redondo  Beach,  California,  south 
to  Punta  Eugenia,  Baja  California. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


209 


B 


FIGURE  12.  Anterior  part  of  body  of  four  species  of  eastern  Pacific  Syngnathidae.  (A)  Syngnathus  euchrous,  holotype,  SIO 
H47-68.  (B)  Syngnathus  euchrous  ollotropis,  holotype,  SIO  H48-53.  (C)  Syngnathus  leptorhynchus,  a  166-mm-SL  male,  SIO 
63-1053.  (D)  Syngnathus  insulae,  holotype,  SIO  63-169. 


MATERIAL  EXAMINED.— Holotype:  SIO  H47-68,  a  232-mm- 
SL  mature  male  collected  at  Bahi'a  Todos  Santos,  Baja  Cali- 
fornia (31°52'N,  116°38'W),  with  a  25-ft  (7.6-m)  beach  seine 
at  a  depth  of  0.3-1.5  m  on  22  May  1947,  by  Carl  L.  and  Laura 
C.  Hubbs. 

Paratypes:  SIO  H47-68,  86(83-228),  same  data  as  holotype. 
SIO  H48-319,  16(95-204),  E  side  of  southern  Isla  Coronado, 
Baja  California,  dredge,  11  m,  22  Dec.  1948,  by  Clark  Hubbs 
and  party  aboard  M/V  ORCA.  GCRL  15757,  22(96-175), 
Scripps  Pier,  La  Jolla,  California,  dip  net  under  night  light,  17 
Aug.  1947,  by  J.  L.  McHugh.  CAS  27405,  11(39-186),  Estero 
de  Punta  Banda,  Baja  California,  8  Sep.  1949,  by  Chaffy, 
Scott,  and  Mead. 

ADDITIONAL  MATERIAL. — California:  Redondo  Pier,  SIO 
H48-204,  1(225).  Scripps  Pier,  SIO  H46-63,  20(54-179).  La 
Jolla  Shores,  SIO  H50-214,  2(175-249);  SIO  H50-227,  5(119- 
139). 

Baja  California:  Islas  Coronados,  SIO  H49-217,  17(142- 
224);  SIO  H49-19,  15(152-223);  SIO  H46-125,  4(86-118);  SIO 


H46-119,  11(132-222);  SIO  H48-217,  14(107-238).  Bahia  de 
Todos  Santos,  SIO  H45-223a,  4(111-213);  SIO  H51-48,  5(99- 
130).  Bahi'a  Blanca,  SIO  60-327,  5(162-203).  Isla  Cedros,  SIO 
53-110,  6(123-186);  SIO  68-280,  1(113). 

INTERGRADES  with   5.   e.  ollotropis:   Punta  Malarrimo, 
UCLA  W51-223,  6(61-223);  UCLA  W5 1-221,  12(76-149). 


Syngnathus  euchrous  ollotropis  n.subsp. 

(Figure  12B) 

DIAGNOSIS. — An  S.  euchrous  with  33-40  (36- 
38)  dorsal  rays;  11-13  pectoral  rays;  3  anal  rays; 
18-20  (19)  trunk  rings;  41-45  (43)  tail  rings;  60- 
65  total  rings;  snout  ridge  well  developed  from 
vomer  to  anterior  third  of  orbit,  rugose,  typically 
without  distinct  rise  anterior  to  orbit;  maximum 
known  size  224  mm  SL;  smallest  sexually  ma- 


210 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


ture  male  102  mm  SL.  See  Tables  5  and  6  for 
additional  counts  and  measurements. 

DESCRIPTION. — Young.  14-16  mm  SL  at 
birth.  All  fins  developed  and  with  rays.  Rings 
without  prominent  spines  but  distinct  triangular 
ridges.  Uniform  orange-tan  with  middle  rays  of 
caudal  fin  more  darkly  pigmented  than  outer. 
Dorsal  and  pectoral  fins  colorless.  Anal  fin  dark- 
ly pigmented. 

Coloration  in  life.  From  C.  L.  Hubbs's  field 
notes  for  SIO  H48-53: 

The  lower  part  of  opercles  dark  and  mottled  ....  Streaked 
color  phase  frequent  and  light  bars  usually  faint  and  seldom 
well  developed  .  .  .  the  lower  parts  are  almost  always 
marked  with  dark  and  usually  variegated  rather  than  being 
clear.  In  general  the  color  pattern  is  more  longitudinal  and 
less  transverse.  In  some  specimens  the  lower  keel  carries 
a  series  of  light  blotches  one  per  segment  or  may  be  largely 
clear.  The  general  color  varies  from  light  tan  to  very  deep 
purple-brown,  almost  black.  The  prominent  marking  of  the 
lower  surface  is  perhaps  the  most  outstanding  color  feature. 

Coloration  in  alcohol.  Typically  brownish 
above  and  light  below,  becoming  entirely 
brownish  on  posterior  third  of  body.  Dorsal  fin 
hyaline.  Anal  and  pectoral  rays  bordered  by 
melanophores.  Often  a  dark  midventral  stripe  on 
trunk.  A  chocolate-brown  band  on  trunk  be- 
tween lateral  trunk  ridge  and  superior  tail  ridge. 
Caudal  with  light  and  dark  reticulations.  A  re- 
ticulated area  around  orbit  and  on  snout.  Most 
often  an  orange-brown  body  with  longitudinal 
lighter  streaks. 

HABITAT. — Eelgrass  beds  and  drifting  algae. 

COMPARISONS. — S.  e.  euchrous  is  easily  dis- 
tinguishable from  5.  e.  ollotropis  by  the  char- 
acters given  in  the  key.  There  is,  however,  a 
zone  of  intergradation  along  the  coast  near  Pun- 
ta  Malarrimo.  The  specimens  examined  from 
there  are  intermediate  in  meristic  characters  be- 
tween the  two  subspecies  (Table  6).  Although 
these  intergrades  are  closer  to  S.  e.  euchrous  in 
the  ranges  of  these  meristic  characters,  they 
have  the  distinctive  straight  snout  ridge  char- 
acteristic of  S.  e.  ollotropis. 

The  semi-isolation  of  S.  e.  ollotropis  in  Turtle 
Bay  has  produced  a  distinctive  morphology 
which  would  have  to  be  considered  specifically 
distinct  had  not  the  intergrades  been  collected. 

The  existence  of  definite  intergrades  between 
S.  e.  euchrous  and  S.  e.  ollotropis  is  concrete 
indication  that  subspecific  recognition  is  war- 
ranted. 


RANGE. — Bahia  Tortugas  south  to  Bahia  de 
Ballenas,  and  Isla  Cedros,  Baja  California. 

MATERIAL  EXAMINED. — Holotype:  SIO  H48-53,  a  224-mm- 
SL  mature  male  collected  %  mile  [1.2  km]  E  of  Cabo  Tortolo, 
Bahia  Tortugas,  Baja  California  (27°38'53"N,  114°51'46"W), 
from  eelgrass  at  a  depth  of  1  m,  with  an  8-ft  [2.4-m]  minnow 
seine,  8  Feb.  1948,  by  Clark  and  Earl  L.  Hubbs. 

Paratypes:  All  collected  in  Baja  California.  SIO  H48-53, 
35(43-220),  same  data  as  holotype.  SIO  H48-55,  47(66-216), 
same  locality  as  holotype,  1.5  m,  25-ft  [7.6-m]  bag  seine,  9 
Feb.  1948,  by  Clark  and  Earl  Hubbs.  GCRL  15758,  15(61- 
158),  same  locality,  1  m,  25-ft  [7.6  m]  bag  seine,  7  Feb.  1948, 
by  Clark  and  Earl  Hubbs.  SIO  H47-112,  1(207),  Bahia  Tor- 
tugas, from  drifting  brown  algae,  23  May  1947,  by  Robert  S. 
Arthur.  SIO  54-131,  1(139),  SE  part  of  Bahia  Tortugas,  shell 
dredge,  R/V  ORCA,  18  July  1954,  by  J.  W.  Sefton,  Jr.  SU 
19260,  1(144),  off  Isla  Cedros,  27  Mar.  1952,  by  James  E. 
Bohlke  and  party,  Sefton-Stanford  ORCA  Expedition. 

ADDITIONAL  MATERIAL. — Bahia  Tortugas,  CAS  5438-5440, 
3(104-112).  Bahfade  Ballenas,  SIO  64-847,  1(135). 

Syngnathus  leptorhynchus  Girard 

(Figures  12C,  13,  14) 

Syngnathus  brevirostris  GIRARD,  1854:156  (original  descrip- 
tion; San  Diego,  California;  cotypes  USNM  969;  nee  S. 
brevirostris  Riippell,  1840). 

Syngnathus  leptorhynchus  GIRARD,  1854  (Oct.  6):  156  (original 
description;  San  Diego,  California;  holotype  USNM  970, 
lost);  JORDAN  AND  EVERMANN  1896:764  (description; 
range;  synonymy);  JORDAN  AND  GILBERT  1880:23  (synon- 
ymy); STARKS  AND  MORRIS  1907: 184  (habits;  range);  HUBBS 
1916:158  (description);  1921:1  (synonymy;  description);  UL- 
REY  AND  GREELEY  1928:42  (synonymy;  range);  ULREY 
1929:6  (listed);  JORDAN,  EVERMANN,  AND  CLARK  1930:240 
(synonymy:  range):  BARNHART  1936:34  (description);  MIL- 
LER AND  LEA  1972:87  (description;  range). 

Syngnathus  californiensis  leptorhynchus;  HERALD  1940:64 
(new  combination;  diagnosed  in  key):  1941:66  (description: 
range;  synonymy). 

Siphostoma  leptorhynchus;  JORDAN  AND  GILBERT  1881:453 
(synonymy);  JORDAN  AND  JOUY  1881:1  (5.  dimidiatus  as 
synonym);  JORDAN  AND  GILBERT  1882:69  (range);  SWAIN 
AND  MEEK  1884:238  (key);  JORDAN  AND  GILBERT  1883:384 
(description;  range;  synonymy). 

Siphostoma  leptorhynchum;  SWAIN  1882:311  (synonymy;  de- 
scription). 

Syngnathus  griseolineatus  AYRES,  1854  (Oct.  27):  14  (original 
description:  probably  near  San  Francisco,  California;  ho- 
lotype ANSP  835);  STARKS  1911:177  (breeding  habits); 
HUBBS  1916:159  (measurements;  compared  with  5.  barbar- 
ae);  1921:1  (counts  and  measurements);  JORDAN,  EVER- 
MANN, AND  CLARK  1930:240  (range). 

Syngnathus  californiensis  griseolineatus;  HERALD  1940:64 
(diagnosed  in  key):  1941:65  (description;  range;  synonymy). 

Siphostoma  griseolineatum;  SWAIN  1882:308,  310,  315  (key; 
synonymy;  description);  SWAIN  AND  MEEK  1884:238  (key); 
K.ERMODE  1909:83  (British  Columbia);  EVERMANN  AND  LA- 
TIMER  1910:136  (Tomales  Bay,  California);  KERMODE 
1929:24  (Victoria,  British  Columbia);  1930:22  (Longborough 
Inlet,  British  Columbia);  JORDAN  AND  EVERMANN  1896:764 
(description;  range;  synonymy). 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


211 


Syngnathus  abbotti  GIRARD,   1859:346  (original  description; 

San  Francisco,  California;  holotype  USNM  971). 
Syngnathus  artindinaceus  GIRARD,  1859:346  (original  descrip- 
tion; coast  of  California;  holotype  USNM  972);  JORDAN, 

EVERMANN,  AND  CLARK,  1930:240  (range). 
Syngnathus  dimidiatus  GILL,  1863:284  (replacement  name  for 

Syngnathus  brevirostris  Girard,  preoccupied). 
Siphostoma  bairdianum  (not  of  Dumeril)  SWAIN,   1882:311 

(description;  Santa  Barbara,  California). 
Siphostoma  barbarae  SWAIN  in  SWAIN  AND  MEEK,  1884:238 

(based  on  specimen  described  by  Swain  (1882)  as  5.  bair- 
•  dianum;  holotype  USNM  31253):  JORDAN  AND  EVERMANN 

1896:765  (description;  synonymy;  range). 
Syngnathus  barbarae:  STARKS  AND  MORRIS  1907:185  (Santa 

Barbara);  HUBBS  1916: 159  (description);  ULREY  AND  GREE- 

LEY  1928:42  (synonymy;  range);  JORDAN,  EVERMANN,  AND 

CLARK  1930:240  (range). 
Syngnathus  californiensis  (in  part,  not  of  Storer):  FOWLER 

1921:443  (description);  STARKS  1911:177  (Puget  Sound). 

DIAGNOSIS. — A  Syngnathus  with  28-43  dor- 
sal rays;  11-13  pectoral  rays;  2-3  anal  rays;  16- 
21  trunk  rings;  36—46  tail  rings;  53-63  total  rings; 
dorsal  on  0-1.5  +  6-9  rings;  brood  pouch  cov- 
ering 18-26  rings;  head  5.6-9.3  in  SL;  dorsal  fin 
base  0.9-1.3  in  head;  snout  1.6-2.4  in  head; 
maximum  known  size  299  mm  SL;  smallest  ma- 
ture male  80  mm  SL.  See  Tables  4,  5,  7,  and  8 
for  additional  counts  and  measurements. 

DESCRIPTION. — All  ridges  of  head  and  body 
moderately  developed,  rounded,  and  rather 
smooth  to  slightly  rugose.  Nuchal  and  prenuchal 
ridges  low.  Supraoccipital  ridge  obsolete  and  not 
continuous  with  nuchal  ridge.  Opercular  ridge 
extends  posteriorly  about  one-third  length  of 
opercle.  Snout  ridge  smooth,  low,  with  small 
hump  over  nares.  Superior  ridge  of  pectoral  cov- 
er plate  smooth,  curved;  inferior  ridge  obsoles- 
cent to  absent.  Supraorbital  ridges  smooth. 
Northern  populations  tend  to  have  more  obso- 
lescent ridges.  Trunk  and  tail  ridges  slightly  ru- 
gose. Superior  tail  and  lateral  trunk  ridges  usu- 
ally not  overlapping;  when  overlapping,  only  for 
length  of  last  trunk  ring.  Scutella  of  medium 
size,  subequal  to  half  depth  of  adjoining  plates. 
Lateral  line  consisting  of  6-9  papillae  per  plate. 
No  dermal  flaps. 

Protecting  plates  of  brood  pouch  moderately 
developed  and  as  much  as  two-thirds  as  deep  as 
attached  tail  ring;  edges  of  brood  pouch  folds 
entire.  Eggs  1.3-1.5  mm  in  greatest  diameter, 
arranged  in  1-2  layers  4  across.  Males  with  eggs 
and  young  collected  year  round. 

Fins  moderately  developed.  Caudal  as  long  as 
last  three  tail  rings.  Pectoral  fin  extends  poste- 
riorly over  about  \l/3  rings. 


4)     CJ 

O.    O 


f>     <N     — 


r«i   Ov    OC    (N 


—       I      (N       I     •<*• 


2,   o    o 


^    J  U 


£    g  c    ==  5 


— 
CO  U 


I  I 


212 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


TABLE  7.     MERISTIC  VARIATION  IN  Syngnathus  leptorhynchus. 


Trunk  rings 

Tail  rings 

Locality                    16 

17 

18 

19 

20     21     36     37      38         39         40      41       42        43       44      45 

46 

Vancouver  Is. 

— 

11 

17 

388           42     — 



Puget  Sound 

5 

21 

4 

2      10       12           7       —     — 

— 

Coos  Bay 

1 

51 

9 

1     —     —     —       —       —           —       4       12         14       27       3 

— 

Humboldt  Bay 

3 

64 

19 

1     —                                               —       4       28         28       21       6 

— 

Tomales  Bay 

2 

31 

4 

7      11        16           3 

— 

San  Francisco  Bay 

2 

10 

190* 

52       1     —     —       —       —           —       3       32*      109      107     20 

1 

Morro  Bay 

1 

35 

3 

1     —       1        1        10       15           10       3       —         — 

— 

Pt.  Mugu  Lagoon 

— 

14 

— 

14             63       — 

— 

C       to   Doi-Kot-o 

if 

oama  oaroara 
Newport  Bay 

2 

1  T 

17 

11 

—     —     —     —         1         2             75       11           3       —     — 

_ 

Mission  Bay                        3 

91 

144 

9 

__       4     19       61       74           64     21         3         —       —     — 

— 

San  Diego  Bay                      1 

14** 

9** 

1 

2       12         9**         2     —       — 

— 

Estero  de  Punta  Banda        2 

12 

5 

— 

4         g          4           i          i          

— 

Bahia  San  Quintin 

10 

29 

— 

—     —       61415         3              1     —       —         ___ 

— 

Scammon  Lagoon                3 

3 

1 

— 

—     —       1          1          1             22—         — 

— 

Total                                      9 

146 

443 

267 

55       1      11     41      109      112          104     75      123        168      157     29 

1 

X 

=  18.2  ± 

0.8 

jc  =  41.2  ±  2.3 

Dorsal  rays 

Locality                     28 

29 

30       31 

32 

33       34         35          36        37         38         39       40       41       42 

43 

Vancouver  Is. 







21             6473111 



Puget  Sound 

— 

—       — 

— 

23             87422       —       — 

— 

Coos  Bay 

— 

—       — 

— 

5           4          16          14        10         8         2 

— 

Humboldt  Bay 

— 

—       — 

— 

7           9          12          17        14        12         8 

2 

Tomales  Bay 

— 

—       — 

— 

1             4           9         11           3         4          1          2 

— 

San  Francisco  Bay 

— 

—       — 

— 

19           31         55         61*       60       25         8          1 

— 

Morro  Bay 

— 

—       — 

3 

5        10       10             65         — 

— 

Pt.  Mugu  Lagoon 

— 

—       — 

2 

225             21         —         — 

— 

Santa  Barbara 

— 

—       — 

— 

It       —         —         —       —       —       — 

— 

Newport  Bay 

— 

—       — 

3 

566             7           3           i         _       _       _       _ 

— 

Mission  Bay                         2 

15 

26       38 

53 

51        37        18             6           !_____ 

— 

San  Diego  Bay 

— 

5         3 

8 

621**—         —         —         —       —       —       — 

— 

Estero  de  Punta  Banda       2 

2 

7         5 

3 

___          —         ______ 

— 

Bahia  San  Quintin 

— 

—         4 

14 

783             2         —           !____ 

— 

Scammon  Lagoon 

1 

1       — 

1 

3          1 

— 

Total                                    4 

18 

39       50 

87 

76       73       58           85         98        113         99       56        30        14 

2 

x  =  35.8  ±  3.3 

*  Holotype  of  S.  barbarae. 
t  Holotype  of  5.  abbotti. 
**  Syntypes  of  S.  brevirostris. 


Coloration  in  life.  Mostly  browns,  purples, 
and  greens,  with  varying  amounts  of  mottling, 
spots,  etc. 

Coloration  in  alcohol.  Coloration  highly  vari- 
able, mainly  shades  of  brown.  Great  inter-  and 
intra-population  variability.  Highly  mottled, 
blotched,  and  striped.  Dorsal  fin  always  with 
some  brownish  spots  at  base  and  fin  rays  bor- 
dered with  melanophores. 


HABITAT. — Ordinarily  inhabiting  eelgrass  in 
bays  and  occasionally  near  shore  outside. 

VARIATION  AND  GROWTH. — Variation:  Syng- 
nathus leptorhynchus  shows  extreme  variation 
in  meristic  characters  (see  Table  7)  approaching 
the  North  Atlantic  species  5.  typhle  in  this  re- 
gard. Duncker  (1908)  statistically  analyzed  sev- 
eral meristic  characters  of  5.  typhle  populations 
from  the  Baltic  Sea,  Plymouth,  and  Naples  and 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


213 


showed  a  definite  clinal  pattern  in  the  variation 
of  numbers  of  trunk  rings  and  pectoral  rays. 
Other  meristic  features  showed  populational  dif- 
ferences, but  in  a  mosaic  pattern  of  different  se- 
quences. 

Meristic  variation  in  S.  leptorhynchus  is  mo- 
saic, and  no  single  character  allows  or  justifies 
recognition  of  more  than  one  species  or  subspe- 
cies. Herald  (1940,  1941)  and  others  have  used 
the  same  meristic  characters  herein  analyzed  in 
support  of  recognizing  northern  and  southern 
species  or  subspecies.  A  mosaic  pattern  of  vari- 
ation is  not  limited  to  Syngnathus  species,  e.g., 
Rosenblatt  (1963)  reported  a  similar  pattern  for 
populations  of  the  Central  American  tripterygiid 
Enneanectes  sexmaculatus. 

Each  bay  along  the  west  coast  of  North  Amer- 
ica appears,  on  the  basis  of  available  evidence, 
to  support  a  distinctive  population  of  S.  lepto- 
rhynchus. Meristic  features  of  each  population 
are  not  stable,  but  do  fluctuate  with  changes  in 
environmental  temperature. 

To  underscore  this  point,  C.  L.  Hubbs  ana- 
lyzed three  collections  from  the  northwest  cor- 
ner of  Mission  Bay  on  22-24  April  1923.  All 
three  collections  were  made  with  exactly  the 
same  method  (seine)  and  at  the  same  location  in 
the  bay.  For  analysis,  all  specimens  97  mm  SL 
or  less  are  considered  to  be  winter  reared,  and 
in  view  of  the  demonstrated  growth  (see  below) 
it  may  be  presumed  that  all  specimens  longer 
than  97  mm  SL  were  at  least  mostly  reared  in 
warmer  water.  The  summary  of  meristic  data 
(Table  8)  compiled  for  these  collections  indi- 
cates that  the  winter-reared  young  have  higher 
means  in  all  meristic  characters  studied  than  do 
the  larger  (adult)  fish.  The  means  for  these  two 
size  groups  are  all  significantly  different  (P  <^ 
.005). 

Although  Mission  Bay  has  been  highly  modi- 
fied in  recent  years,  the  data  are  indicative  of 
changes  in  meristic  characters  as  a  result  of  en- 
vironmental temperature.  Similar  seasonal  me- 
ristic variation  has  been  documented  by  Broth- 
ers (1975)  for  the  goby  Quietula  y-cauda — using 
samples  taken  throughout  the  year,  he  deter- 
mined that  the  mean  vertebral  number  of  the 
postlarvae  is  inversely  correlated  with  seasonal 
changes  in  environmental  temperature. 

Each  local  population  of  Syngnathus  lepto- 
rhynchus is  therefore  distinctive  and  also  highly 
modified  by  local  environmental  conditions.  S. 


VI     VI 

o  o 


c  .r: 

3     3 

o  -a 


214 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


Imm 
FIGURE  13.    Syngnathus  leptorhynchus  at  time  of  emergence  from  brood  pouch.  16.3  mm  SL. 


leptorhynchus  may  be  a  useful  indicator  species 
for  the  effects  of  pollution. 

Growth:  On  16  June  1974,  a  male  5.  lepto- 
rhynchus (165  mm  SL)  gave  birth  to  85  young 
in  the  laboratory  of  the  National  Marine  Fish- 
eries Service,  La  Jolla.  The  young  were  imme- 
diately separated  from  their  father  and  kept  alive 
until  a  failure  in  the  seawater  system  on  6  Sep. 
The  young  were  fed  Anemia  nauplii  and  fresh- 
caught  plankton.  An  overabundance  of  food  was 
always  present.  Samples  of  the  young  were  re- 
moved frequently  during  the  period  from  16  June 
to  6  Sep.  and  preserved  in  10%  formalin  for  later 
analysis. 

Figure  14  represents  the  measured  growth  of 
the  young  for  the  period.  The  data  have  been 
analyzed  and  fitted  to  a  Laird-Gompertz  equa- 
tion (Laird,  Barton,  and  Tyler  1968)  calculated 
after  the  methods  of  Zweifel  and  Lasker  (1976). 
The  Laird-Gompertz  equation  is  preferred  when 
analyzing  growth  for  periods  less  than  one  year. 

On  16  Aug.  development  of  the  brood  pouch 
had  already  begun  in  a  76-mm-SL  individual. 
Four  males  (80-93  mm  SL),  each  with  a  fully 
developed  brood  pouch,  were  found  on  24  Aug. 
By  the  termination  of  the  experiment  on  6  Sep., 
the  largest  specimen  was  a  fully-mature,  1 14-mm 
male. 

That  the  ranges  of  meristic  variation  for  the 
adults  do  not  completely  subtend  the  ranges  for 
the  young  together  with  rapid  growth  to  sexual 
maturity  strongly  indicate  that  individuals  of  5. 
leptorhynchus  (at  least  from  Mission  Bay)  prob- 
ably live  for  only  one  year.  This  is  in  agreement 
with  the  data  presented  by  Brown  (1972)  for  S. 
louisianae,  S.  scovelli,  S.  floridae,  and  Micro- 
gnathus  crinigerus  for  Cedar  Key,  Florida.  Based 
on  length-frequency  analyses,  Brown  estimated 
a  life  span  of  perhaps  one  year. 

COMPARISONS. — A  highly  variable  species 
that  is  similar  to  the  Japanese  S.  schlegeli.  In 
fact,  it  is  rather  difficult  to  separate  members  of 
northern  populations  of  leptorhynchus  from 


those  of  schlegeli.  However,  S.  schlegeli  has  a 
longer  snout  and  longer  fins,  and  the  ridges  of 
the  body  are  better  developed.  S.  insulae,  also 
closely  related  to  S.  leptorhynchus,  may  be  dis- 
tinguished by  the  characters  given  in  the  account 
of  that  species. 

RANGE. — Southeastern  Alaska  south  to  Bahfa 
Santa  Maria,  southern  Baja  California. 

MATERIAL  EXAMINED.— Alaska:  UW  14486,  1(179);  UW 
1610,  1(212). 

British  Columbia:  Vancouver,  BC  53-253a,  28(60-206);  SIO 
62-206,  2(137-192). 

Washington:  Puget  Sound,  CAS  10935,  3(125-140);  UW 
4927,  3(136-151);  UW  52%,  2(146-245);  UW  4258,  3(86-130); 
UW  4918,  4(139-167);  SIO  74-73,  19(73-215). 

Oregon:  Tillamook,  UMMZ  93951,  33(49-235).  Coos  Bay, 
UMMZ  93956,  63(35-235). 

California:  Klamath  Lagoon,  CAS  45070,  1(216).  Humboldt 
Bay,  HSU  OT62-7,  44(136-207);  HSU  OT61-18,  14(121-218); 
HSU  OT64-5,  12(110-210).  Mendocino  County,  SU  20515, 
5(165-214).  Tomales  Bay,  SIO  75-459,  37(54-234);  UCLA 
W55-198,  35(61-207);  CAS  19860,  2(211-293);  CAS  26207, 
2(104-252).  Drakes  Bay,  CAS  7518,  1(130).  San  Pablo  Bay, 
CAS  45072,  33(59-272).  San  Francisco  Bay,  SU  4190,  24(115- 
159);  USNM  971,  1(228),  holotype  of  S.  arundinaceus; 
UMMZ  64416,  1(215);  UMMZ  6044,  1(151);  CAS  ALBATROSS 
Biological  Survey  of  San  Francisco  Bay*,  D5826,  1(131); 
H5159,  2(139-165);  D5816,  1(129);  H5311,  1(119);  H5013, 
4(111-184);  D5847,  1(92);  H5003,  1(87-121);  H5318,  2(110- 
128);  H5185,  1(61);  D5848,  2(101-118);  H5312,  2(66-94): 
H5004,  2(77-110);  H5308,  1(67);  H5230,  1(62):  H5248,  1(64): 
H5259,  1(61);  H5310,  1(80):  H5316,  2(69-98):  H5309,  2(106- 
120):  H5250,  7(51-123);  H5009,  9(33-109):  H5257,  1(86); 
H5261,  2(41-43);  H5345,  13(25-133);  H5313,  6(69-93);  H5251, 
6(73-125);  H5010,  2(33-66);  H5005,  3(44-69);  H5275,  3(38- 
70);  H5315,  5(63-113);  H5252,  5(62-95):  H5317,  4(76-103); 
H5311,  8(75-90):  H5274,  8(67-77);  H5007,  11(28-108):  H5314, 
13(67-105);  H5272,  17(64-146);  H5276,  25(49-130);  H5006, 
76(27-116).  Elkhorn  Slough,  SIO  62-510,  3(77-103).  Del 
Monte,  SIO  48-372,  2(92-196).  Morro  Bay,  SIO  H50-191, 
1(243);  UCLA  W50-196,  50(98-236):  SU  48888,  10(162-217). 
Point  Mugu,  CAS  45076,  from  E.  S.  Herald  personal  collec- 
tion, 4(151-181).  Santa  Barbara,  USNM  31253,  type  of  5. 
barbarae.  San  Pedro,  SIO  H48-180,  1(257).  Newport  Bay, 
UCLA  W50-171,  50(99-278);  UCLA  W50-172,  3(101-132); 
SIO  H46-33,  5(215-280).  San  Juan  Capistrano,  SIO  H52-109, 
1(101).  Mission  Bay,  SIO  H45-8,  50(72-198);  SIO  H46-10, 


Station  data  may  be  found  in  Anonymous  1921. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 
IZOr 


215 


100 


80 


SL 


60 


40 


20 


L,«l5.5e 


2.04(l-e°3t) 


0  10  20  30 

DAYS   AFTER    BIRTH 


40 


50 


60 


70 


80 


FIGURE  14.     Growth  of  Syngnathus  leptorhynchus.  Standard  length  plotted  against  time  in  days.  The  Laird-Gompertz  Equa- 
tion fitted  to  these  data  is  included. 


3(116-125).  San  Diego  Bay,  USNM  969,  3,  types  of  5.  bre- 
virostris;  SIO  64-337,  25(100-158). 

Baja  California:  Estero  de  Punta  Banda,  SIO  H45-15, 
19(104-158).  Bahfa  San  Quintin,  SIO  60-453,  128(69-245);  SIO 
60-450,  6:  SU  15251,  10(123-299).  Laguna  Scammon,  SIO 
H52-140,  1(95).  Bahfa  Santa  Maria,  SIO  H50-149;  SIO  60-294, 
1(94). 

Syngnathus  insulae  n.sp. 

(Figure  12D) 

DIAGNOSIS. — A  Syngnathus  with  31-34  (32- 
33)  dorsal  rays;  10-12  pectoral  rays;  2-4  anal 
rays;  17-19  (18)  trunk  rings;  40-42  (41-42)  tail 
rings;  58-60  total  rings;  dorsal  on  0.2-1 .2  +  6.5- 
8  rings;  19  rings  covered  by  brood  pouch;  head 
8.6-9.3  in  SL;  dorsal  fin  base  1.0  in  head;  snout 
1.7-2.3  in  head;  maximum  known  size  204  mm 
SL;  smallest  sexually  mature  male  125  mm  SL. 
See  Tables  4  and  5  for  additional  measurements. 

DESCRIPTION. — All  ridges  of  head  and  body 
poorly  developed,  smooth.  Prenuchal  and  nu- 
chal crests  obsolete.  Opercular  ridge  reduced  to 
short  basal  portion.  Ridges  of  pectoral  cover 
plate  low.  Trunk  and  tail  ridges  low,  rounded 
but  distinguishable.  Superior  tail  ridge  not 
reaching  beyond  first  tail  ring,  only  overlapping 
lateral  trunk  ridge  for  one-half  ring.  Scutella 
quite  distinct  and  about  half  as  deep  as  adjoining 
plate.  Lateral  line  consisting  of  5-6  papillae  per 
plate.  No  dermal  flaps. 


Protecting  plates  of  brood  pouch  well  devel- 
oped, each  about  as  deep  as  adjacent  tail  ring; 
edges  of  brood  pouch  folds  entire.  Eggs  1.2-1.3 
mm  in  greatest  diameter,  arranged  in  one  layer 
two  across.  Males  with  eggs  and  young  have 
been  collected  in  April. 

Fins  normally  developed.  Caudal  fin  as  long 
as  last  three  tail  rings.  Pectoral  fin  reaches  pos- 
teriorly to  end  of  first  trunk  ring. 

Young.  About  12  mm  SL  at  birth.  About  five 
dark  bands  posterior  to  vent.  Body  sprinkled 
with  small  round  melanophores,  especially  in 
the  area  of  the  bands.  Snout  short,  mouth  dis- 
tinctly upturned. 

Coloration  in  life.  From  C.  L.  Hubbs's  field 
notes  for  SIO  H46- 143: 

The  greener  specimen  has  a  dark  streak  running  downward 
and  backward  from  eye.  In  this  specimen  the  streak  ex- 
tending upward  and  backward  from  eye  is  less  distinct.  The 
streak  running  forward  from  eye  to  chin  is  conspicuous  pos- 
teriorly but  interrupted  anteriorly.  The  redder  specimen  has 
a  dark  spot  just  behind  and  below  eye,  but  no  definite 
streaks  radiating  from  eye. 

Coloration  in  alcohol.  A  light  tan  to  tan  fish 
without  any  distinguishing  pattern.  The  caudal 
fin  may  be  lighter  at  its  border.  Darker  speci- 
mens may  have  a  sprinkling  of  small  melano- 
phores bordering  rays. 

HABITAT. — Occasionally  in  floating  Sargas- 


216 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


sum,  but  much  more  often  taken  at  depths  of 
20-34  m  in  beds  of  Sargassum  and  algae. 

COMPARISONS. — Appears  most  closely  related 
to  5.  leptorhynchus.  The  two  are  very  close  in 
meristic  characters.  S.  leptorhynchus  has  the 
longer  snout  and  postorbital.  5.  insulae  differs 
from  leptorhynchus  in  the  broader  interorbital, 
smaller  orbit,  and  less  robust  body. 

RANGE. — Endemic  to  Isla  Guadalupe,  an 
oceanic  island  well  off  the  coast  of  Baja  Cali- 
fornia. 

ETYMOLOGY. — From  the  Latin  insulae,  gen- 
itive of  insula  (island). 

MATERIAL  EXAMINED. — Holotype:  SIO  63-169,  a  160-mm- 
SL  mature  male  collected  at  Caleta  Malpomene,  a  cove  at  the 
south  end  of  Isla  Guadalupe,  Mexico,  from  a  depth  of  22-50 
m  using  a  try-net  on  22  Apr.  1963,  by  C.  L.  Hubbs  and  party 
aboard  the  R/V  HORIZON. 

Paratypes:  All  taken  from  Isla  Guadalupe.  SIO  63-169, 
32(77-204),  same  data  as  holotype.  SIO  H46-143,  2(98-98.4), 
Caleta  Melpomene,  under  nightlight,  6  Dec.  1946,  by  C.  L. 
Hubbs  aboard  R/V  E.  W.  SCRIPPS.  SIO  H46-144,  1(118),  Cal- 
eta Melpomene,  3-6  m,  dredge,  7  Dec.  1946,  by  C.  L.  Hubbs 
aboard  R/V  E.  W.  SCRIPPS.  GCRL  15756,  2(93-117),  Caleta 
Melpomene,  under  nightlight,  27  Jan.  1950,  by  C.  L.  Hubbs 
and  party  aboard  R/V  ORCA.  SIO  H50-31,  1(99),  Caleta  Mel- 
pomene, 3-7  m,  dredge,  29  Jan.  1950,  by  C.  L.  Hubbs  and 
party  aboard  R/V  ORCA.  SIO  54-214,  2(51-87),  '/s  mile  [200  m] 
"off  Barracks"  (Northeast  Anchorage),  under  nightlight,  13 
Nov.  1964,  byC.  L.  Hubbs  and  party.  SIO  63-188,  1(1 16),  just 
inside  s  side  Punta  Amarillo  on  rocky  bottom,  15  m,  Chemfish 
and  SCUBA,  26  Apr.  1963,  by  J.  R.  Stewart  and  party. 

FOSSIL  SYNGNATHIDAE  OF  SOUTHERN 
CALIFORNIA 

The  following  discussion  of  the  fossil  pipe- 
fishes of  southern  California  is  based  on  a  large 
collection  in  the  Section  of  Vertebrate  Paleon- 
tology at  the  Natural  History  Museum  of  Los 
Angeles  County  (LACM),  including  the  collec- 
tions formerly  housed  at  the  California  Institute 
of  Technology  (CIT). 

The  fossil  syngnathids  from  southern  Califor- 
nia are  all  of  Miocene  to  possibly  Pliocene  age. 
Most  specimens  have  been  collected  in  diato- 
mites  and  diatomaceous  siltstones  of  the  upper 
Modelo  and  Puente  formations,  although  a  num- 
ber have  been  collected  from  argillaceous  diat- 
omites  of  the  lower  Modelo  Formation.  Locali- 
ties are  given  by  number,  and  more  precise  data 
can  be  obtained  by  referring  to  the  Locality 
Catalogue  in  the  Vertebrate  Paleontology  Sec- 
tion (LACM). 

These  formations  have  provided  the  bulk  of 
material  used  in  the  descriptions  of  California 


fossil  fish  (Jordan  and  Gilbert  1919;  David  1943). 
Since  most  of  the  fishes  contained  in  these  de- 
posits are  pelagic  to  bathypelagic,  previous  au- 
thors have  inferred  that  during  the  Miocene  this 
area  was  a  large  basin  with  depths  ranging  from 
200-500  m  (David  1943)  to  1000  m  (Crane  1966). 

Materials  and  Methods 

The  anal  and  pectoral  fins  of  fossil  pipefishes 
are  usually  not  visible.  Since  the  anal  fin  marks 
the  first  tail  ring,  it  is  impossible  to  make  precise 
counts  of  trunk  and  tail  rings.  However,  the 
number  of  plates  in  a  ring  around  the  trunk  is 
usually  seven  and  there  are  four  around  the  tail. 
When  these  plates  could  be  seen,  accurate 
counts  were  made.  When  such  counts  were  not 
determinable,  the  trunk  rings  were  counted  as 
ending  at  the  first  ring  under  the  dorsal  fin  or  the 
posterior  end  of  the  median  ventral  ridge  (in 
Hipposyngnathus). 

The  count  of  dorsal  fin  rays  in  fossil  pipefishes 
is  difficult  because  of  frequent  splitting  of  the 
two  ray-halves.  However,  with  proper  precau- 
tion, a  fairly  accurate  count  is  often  possible. 

Accurate  counts  of  the  rings  and  dorsal  rays 
are  of  particular  importance  in  syngnathid  tax- 
onomy, as  these  counts  provide  the  primary  ba- 
sis for  classification  of  living  forms. 

Correct  generic  allotment  of  fossil  syngnathid 
species  is  based  primarily  on  the  author's 
knowledge  of  the  general  appearance  of  syng- 
nathid genera.  The  characters  often  used  for 
placement  of  syngnathids  into  a  genus  (i.e.,  lat- 
eral ridge  pattern,  placement  and  position  of 
brood  pouch)  are  very  rarely  preserved  in  fos- 
silized pipefishes. 

This  section  on  fossil  pipefishes  is  intended 
only  to  describe  those  forms  known  to  occur  in 
deposits  along  the  eastern  Pacific  and  is  not 
meant  to  be  a  critical  review  or  stratigraphic 
analysis  of  them. 

Subfamily  Hipposyngnathinae  nom.  nov. 

Eogastrophinae  JERZMANSKA  1968:436  ("genre  typique:  Hip- 
posyngnathus  Danil'chenko  1960"). 

Since  the  subfamily  name  was  incorrectly  pro- 
posed as  Eogastrophinae  (the  sole-included  ge- 
nus is  Hipposyngnathus),  and  since  subfamilial 
ranking  is  obviously  called  for,  I  propose  the 
name  Hipposyngnathinae,  with  Hipposyngna- 
thus  as  the  type-genus. 

Jerzmanka  diagnosed  the  subfamily  adequate- 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


217 


B 


FIGURE  15.     Holotypes  of  three  species  of  California  fossil  syngnathids.  (A)  Hipposyngnathus  imporcitor,  LACM  12055. 
(B)  Syngnathus  avus,  CAS  Geology  Dept.  No.  58372.  (C)  Syngnathus  emeritus,  LACM  27445. 


ly  except  that  she  incorrectly  stated  that  scutella 
are  absent. 

Hipposyngnathus  Danil'chenko 

Hipposyngnathus  DANIL'CHENKO,  1960:87  (type-species  by 
original  designation  H.  convexus  Danil'chenko);  JERZMAN- 
SKA  1968:436. 

DIAGNOSIS. — Lateral  protecting  plates  of 
brood  pouch  absent,  however,  a  very  pro- 
nounced median  ventral  trunk  ridge  probably  in- 
dicates position  of  brood  area.  Dorsal,  pectoral, 
and  caudal  fins  present.  Dorsal  mainly  on  the 
tail.  Ridges  of  body  generally  moderately  de- 
veloped and  rugose,  but  without  spines  or  ser- 
rations. Three  described  species  from  the  Oli- 
gocene  and  Miocene. 

DISTRIBUTION. — Oligocene  of  the  Caucasus 
and  Carpathian  Mountains  of  southeastern  Eu- 
rope. Miocene  of  southern  California. 

Hipposyngnathus  imporcitor  n.sp. 

(Figure  15A) 

Syngnathus  avus  (in  part,  not  of  Jordan  and  Gilbert,  1919): 
DAVID  1943:70  (description). 


Holotype:  LACM  12055,  an  almost-complete  specimen  from 
the  upper  Modelo  Formation  of  the  northeast  slope  of  Santa 
Monica  Mountains,  Los  Angeles  County,  California  (Locality 
LACM  (CIT)  326). 

REFERRED  MATERIAL  (all  from  California). — Upper  Modelo 
Formation,  NE  slope  Santa  Monica  Mountains,  Los  Angeles 
County,  24  identifiable  specimens,  33  fragments,  LACM  loc. 
no.  (CIT)326:  LACM  cat.  nos.  10060-10061,  10063,  10114, 
10134,  10143,  10145,  12054,  25341-25351;  and  LACM  loc.  no. 
1267:  LACM  cat.  nos.  13362-13373,  13451-13465,  13469- 
13473,  13475,  13477-13478,  25789-25791.  Near  Lompoc, 
LACM  loc.  no.  6589,  10  fragmentary  specimens:  LACM  cat. 
nos.  12160-12169.  Near  Emery  Park,  Alhambra,  Los  Angeles 
County,  LACM  loc.  nos.  1031,  1033,  14  specimens:  LACM 
cat.  nos.  1056-1058,  1349,  12459-12464,  25668.  Dixie  Canyon, 
Santa  Barbara  County,  LACM  loc.  no.  (CIT)329,  6  speci- 
mens: LACM  cat.  nos.  25357-25359,  25363-25365. 

DIAGNOSIS. — A  Hipposyngnathus  from  the 
upper  Modelo  Formation  of  southern  California 
with  about  22  dorsal  rays;  20-22  trunk  rings;  42- 
44  tail  rings;  snout  1.5-1.9  in  head;  orbit  7.8- 
12.0  in  head;  pouch  rings  equal  to  trunk  rings; 
pectoral  fin  present  but  rays  not  countable; 
pouch  ridge  as  long  as  adjacent  trunk.  Maximum 
size  known  242  mm  SL. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


DESCRIPTION. — Body  ridges  moderately  de- 
veloped and  rugose.  Plates  rhomboid-ovate  with 
10-13  ridgelets  on  either  side  of  median  ridge, 
large;  each  trunk  plate  about  as  wide  as  orbit. 
Opercular  ridge  smooth,  about  half  as  long  as 
opercle.  Snout  without  distinct  ridge.  Scutella 
small.  Two  infraorbitals.  Vertebrae  as  in  Syng- 
nathus.  Dorsal  high,  about  1.5  times  higher  than 
adjacent  tail.  Caudal  normally  developed. 

DISCUSSION. — H.  imporcitor  is  the  most  com- 
mon syngnathid  in  the  upper  Modelo  Formation. 
The  extinct  Hipposyngnathus  was  previously 
known  from  two  European  Oligocene  species. 
Hipposyngnathus  imporcitor  can  be  easily  dis- 
tinguished from  its  congeners  H.  neriticus  and 
H.  convexus  by  the  characters  given  in  the  di- 
agnosis. 

The  genus  Hipposyngnathus  is  unusual  in 
having  a  very  exaggerated  ventral  trunk  ridge. 
Jerzmanska  (1968)  suggested  that  the  males 
alone  had  this  ridge  and  that  it  was  the  site  of 
egg  deposition.  She  also  speculated  that  pouch 
flaps  were  present  on  either  side  of  the  ridge. 
However,  the  extant  genus  Maroubra  also  has 
an  enlarged  ventral  trunk  ridge,  although  not  so 
well  developed  as  in  Hipposyngnathus,  and  its 
eggs  are  not  protected  by  flaps.  It  seems  im- 
probable that  Hipposyngnathus  had  brood- 
pouch  flaps. 

The  Gastrophori  are  typically  spiny  with  the 
exception  of  Nerophis  and  Entelurus.  Hippo- 
syngnathus is  most  like  Nerophis  in  general 
body  form. 

PALEOECOLOGY. — Crane  (1966)  has  discussed 
the  paleoecology  of  Chauliodus  eximius,  a  chau- 
liodontid  from  the  diatomite  of  the  upper  Mo- 
delo. Most  of  his  specimens  were  from  the  same 
localities  as  those  where  Hipposyngnathus  was 
collected.  The  upper  Modelo  is  approximately 
15  million  years  old  (Crane  1966). 

David  (1943)  discussed  the  paleoecology  and 
climate  of  the  upper  Sisquoc  Formation  and  the 
deposits  at  Lompoc.  She  concluded  that  the 
Lompoc  deposits  were  younger  and  that  the 
fishes  found  there  were  neritic  and  pelagic,  and 
those  of  the  upper  Modelo  more  bathypelagic. 
Her  conclusion  that  the  fish  fauna  of  the  upper 
Modelo  indicated  a  basin  depth  of  200-500  m  is 
probably  in  error.  Recent  evidence  (Crane  1966) 
supports  the  view  that  the  basin  was  probably 
at  least  1000  m  in  depth. 

With  the  above  evidence  in  mind,  it  seems 


most  probable  that  H.  imporcitor  was  a  pelagic 
species,  which  may  have  accompanied  floating 
algae,  much  as  do  the  Recent  species  Syngna- 
thus  pelagicus  and  S.  californiensis.  This  con- 
clusion is  supported  by  the  evidence  presented 
by  Danil'chenko  (1960)  for  the  fishes  of  the  Mai- 
kop basin.  He  showed  that  this  basin  was  anoxic 
below  a  depth  of  300  m,  therefore  H.  convexus 
may  be  assumed  to  be  a  pelagic  species. 

ETYMOLOGY. — From  the  Latin  Imporcitor — 
"a  diety  that  presides  over  the  drawing  of  fur- 
rows," in  reference  to  the  greatly  expanded 
trunk  ridge.  To  be  treated  as  a  noun  in  apposi- 
tion and  as  of  masculine  gender. 

Syngnathus  Linnaeus 
Syngnathus  emeritus  n.sp. 

(Figure  15C) 

Holotype:  LACM  27445,  an  almost  complete  specimen  from 
the  Puente  Formation  near  San  Dimas,  Los  Angeles  County, 
California  (Locality  LACM  7153). 

REFERRED  MATERIAL. — Puente  Formation  near  San  Dimas, 
Los  Angeles  County,  California,  LACM  loc.  nos.  5153,  7046, 
14  specimens  and  fragments:  LACM  26146-26148,  27431- 
27434,  27437-27440,  27446-27448. 

DIAGNOSIS. — A  Syngnathus  with  43-47  dor- 
sal rays;  10  caudal  rays;  pectoral  rays  not  de- 
tectable; 14-15  trunk  rings;  51  tail  rings;  snout 
1.5-1.8  in  head;  orbit  5.8-8.7  in  head;  brood 
pouch  not  seen;  largest  specimen  examined  181 
mm  SL. 

DESCRIPTION. — Ridges  of  body  smooth  and 
generally  not  accentuated.  Osteology  basically 
the  same  as  in  extant  species  of  Syngnathus. 
Plates  small,  width  of  trunk  plate  less  than  orbit 
diameter.  Dorsal  about  as  high  as  depth  of  ad- 
jacent body. 

COMPARISONS. — S.  emeritus  is  unusual  in 
having  15  trunk  rings  combined  with  51  tail 
rings.  These  characters  and  the  number  of  dorsal 
rays  distinguish  it  from  S.  avus. 

The  relationships  of  5.  emeritus  perhaps  lie 
with  the  extant  species  S.  leptorhynchus  and  S. 
auliscus.  It  is  possible  that  S.  emeritus  was  on 
the  ancestral  line  leading  to  one  or  both  of  these 
species. 

ETYMOLOGY. — From  the  Latin  emeritus — 
honorably  retired. 

PALEOECOLOGY. — S.  emeritus  is  found  in  the 
diatomaceous  shale  of  the  Puente  Formation.  Its 
habits  probably  were  much  like  those  of  S.  avus, 
i.e.,  living  in  relatively  shallow  water  among 
marine  vegetation. 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


219 


Syngnathus  avus  (Jordan  and  Gilbert) 

(Figure  \5B) 

Syngnathus  avus  JORDAN  AND  GILBERT,  1919:34  (Miocene 
chalk  near  Titus  Avenue,  Bairdstown,  California,  U.C.  loc. 
no.  3525:  type-specimen  by  original  designation,  SU  XLIV, 
B  of  original  description):  JORDAN  1921:259  (reconstruc- 
tion); DAVID  1943:70  (in  part;  description;  synonymy). 

Syngnathus  californiensis  avus:  HERALD  194 1:69  (description; 
discussion). 

Holotype:  CAS  Geology  Dept.  no.  58372  (SU  no.  XLIV,  B 
of  original  description),  all  fossil  fish  types  and  other  Jordan 
paleoichthyological  material  is  now  in  the  Geology  Depart- 
ment, California  Academy  of  Sciences. 

Paratypes:  CAS  Geology  Dept.  no.  58373  (SU  no.  V,  also 
marked  115,  A  of  original  description);  CAS  Geology  Dept. 
no.  58374  (SU  no.  1 15A,  C  of  original  description). 

REFERRED  MATERIAL. — Lower  Modelo  Formation,  Santa 
Monica  Mountains,  Los  Angeles  County,  California,  LACM 
loc.  no.  1681,  15  specimens:  LACM  cat.  nos.  11598-11605, 
26438-26444. 

DIAGNOSIS. — A  Syngnathus  with  36  dorsal 
rays;  10  caudal  rays;  18  trunk  rings;  43  tail  rings; 
dorsal  on  0.5  +  7.5  rings;  pouch  covering  22 
rings;  head  6.4-7.0  in  SL;  snout  1.7-1.8  in  head; 
dorsal  fin  base  1 .3  in  head;  maximum  known  size 
225  mm  SL. 

DESCRIPTION. — Generally  as  given  by  Jordan 
and  Gilbert  (1919).  Osteology  of  neurocranium 
same  as  that  described  for  extant  species  of 
Syngnathus  (Jungersen  1910).  Opercles  without 
median  ridge. 

DISCUSSION. — The  type  material  is  now  in 
such  poor  condition  that  the  characters  as  given 
by  Jordan  and  Gilbert  (1919)  cannot  accurately 
be  determined.  However,  specimen  No.  58373 
(paratype)  does  have  a  brood  pouch  with  pro- 
tecting plates  developed;  they  are  not  missing  as 
stated  by  Herald  (1941). 

Syngnathus  avus  is  a  rather  common  member 
of  the  fossil  fauna  of  the  argillaceous  diatomites 
of  the  lower  Modelo  Formation.  The  original 
field  notes  indicate  that  one  slab  measuring  4  by 
4  ft  [1.2  x  1.2  m]  contained  42  specimens. 

It  must  be  pointed  out  that  Figure  2  on  Plate 
XXIX  of  Jordan  and  Gilbert  (1919)  is  not  of  the 
type  of  S.  avus.  In  fact,  Jordan  and  Gilbert  mis- 
takenly included  a  figure  of  a  fossil  pipefish  from 
Austria  rather  than  the  correct  figure  of  5.  avus. 
This  mistake  was  pointed  out  by  Duncker  (1923) 
and  was  subsequently  corrected  by  Jordan 
(1926,  1927). 

DISTRIBUTION  AND  RELATIONSHIPS 
The  eastern  Pacific  syngnathids  are  generally 
distributed,  with  some  exceptions,  in  commonly 


recognized  zoogeographic  provinces.  Most  of 
the  pipefishes  are  found  in  bays  and  sloughs 
where  the  water  temperature  may  be  higher  than 
along  the  adjacent  open  coast.  As  a  result  their 
distribution  often  extends  past  the  higher-lati- 
tude, colder- water  boundary  established  for  the 
open-coast  fauna.  This  kind  of  distribution  is 
shown  by  other  bay-dwelling  fishes  such  as  the 
blennies  and  gobies  (distributions  in  Miller  and 
Lea  1972).  For  example,  Hypsoblennius  gentilis 
ranges  from  the  Gulf  of  California  to  Monterey, 
Ilypnus  gilberti  ranges  from  the  Gulf  of  Califor- 
nia to  Tomales  Bay,  Clevelandia  ios  ranges  from 
the  Gulf  of  California  to  British  Columbia,  and 
Quietula  y-cauda  ranges  from  the  Gulf  of  Cali- 
fornia to  Morro  Bay.  Syngnathus  leptorhynchus 
ranges  from  the  outer  coast  of  Baja  California 
to  southern  Alaska.  Thus,  these  bay-dwelling 
fishes  cross  the  Point  Conception  faunal  bound- 
ary (Briggs  1974). 

There  are  more  species  of  Syngnathus  in  the 
Northern  Hemisphere  than  in  the  Southern 
Hemisphere.  The  close  similarity  of  Japanese 
Syngnathus  schlegeli  to  generalized  members  of 
two  distinct  eastern  Pacific  lineages  supports  the 
hypothesis  that  the  eastern  Pacific  Syngnathus 
are  derivable  from  two  separate  invasions  from 
the  northwest.  An  alternative  hypothesis  is  in- 
vasion of  Japan  from  the  eastern  Pacific  fol- 
lowed by  a  reinvasion  of  the  eastern  Pacific. 
However,  the  first  hypothesis  appears  to  be  the 
most  parsimonious.  One  lineage  consists  of  S. 
carinatus,  S.  macrobrachium,  and  5.  folletti. 
The  other  consists  of  S.  leptorhynchus,  S.  eu- 
chrous,  and  S.  insulae.  The  related  5.  califor- 
niensis and  5.  exilis  occupy  a  somewhat  isolated 
position  in  the  genus  and  may  be  the  products 
of  another  radiation. 

The  problem  of  defining  these  lineages  and 
relating  them  to  S.  schlegeli  is  one  of  not  being 
able  to  determine  plesiomorphic  and  apomor- 
phic  character  states  at  the  species  level  within 
the  genus  Syngnathus.  Thus,  overall  similari- 
ties, including  meristic  characters  and  strength 
of  ridge  development,  become  the  most  impor- 
tant criteria  used. 

The  carinatus-macrobrachium-folletti  lineage 
is  characterized  by  a  relatively  long  snout;  large, 
well-developed  fins;  large  orbit;  and  rather  well- 
developed  ridges.  The  presence  of  carinatus  in 
the  upper  Gulf  of  California  and  macrobrachium 
and  folletti  in  South  America  parallels  the  dis- 


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tribution  given  by  Chen  (1975)  for  the  exsul-spi- 
norbis  and  capensis  complexes  of  the  subgenus 
Sebastomus.  Chen  speculated  that  these  lin- 
eages probably  shared  a  common  ancestor  that 
was  able  to  both  invade  the  Gulf  and  cross  the 
tropics  using  the  same  climatic  event.  He  was, 
however,  unable  to  rule  out  multiple  crossings. 
The  lack  of  any  close  relative  of  S.  carinatus 
along  the  outer  coast  of  Baja  California  or  Cal- 
ifornia is  puzzling  and  quite  different  from  the 
Sebastomus  example  given  above.  It  is  possible 
that  the  carinatus-macrobrachium-folletti  lin- 
eage may  have  had  a  southern  origin,  although 
more  evidence  is  needed  in  support  of  this  hy- 
pothesis. 

The  leptorhynchus-euchrous-insulae  lineage 
is  characterized  by  the  medium-length  snout, 
which  is  round  in  cross  section,  and  an  almost 
straight  snout  ridge  which  does  not  rise  much 
anterior  to  the  orbit.  The  restriction  of  this  group 
essentially  to  Baja  California  northward  is  inter- 
preted to  mean  that,  assuming  a  more  northern 
origin,  this  lineage  is  much  more  recent  than  the 
preceding  one. 

It  is  apparent  that  S.  leptorhynchus  has  given 
rise  to  the  Guadalupe  Island  endemic  5.  insulae, 
probably  as  a  result  of  waif  dispersal  via  the 
California  Current. 

Syngnathus  leptorhynchus  and  S.  euchrous 
probably  share  a  common  ancestor.  The  isola- 
tion of  a  population  of  S.  euchrous  in  Turtle  Bay 
has  produced  the  subspecies  5.  e.  ollotropis. 
Turtle  Bay  is  unusually  warm  and  the  Macro- 
cystis  pyrifera  found  there  has  a  different  growth 
form  than  that  along  the  coast  (Neushul  1971). 
A  zone  of  intergradation  between  5.  e.  euchrous 
and  S.  e.  ollotropis  exists  at  Puerto  Malarrimo. 

Syngnathus  exilis  and  S.  californiensis  are 
characterized  by  an  elongate,  compressed 
snout,  and  rounded  ridges.  It  is  possible  that 
they  are  related  to  the  carinatus-macrobrach- 
ium-folletti  lineage,  but  their  relationships  can- 
not be  determined  at  this  time. 

The  relationships  of  the  fossil  5.  avus  and  5. 
emeritus  are  difficult  to  determine.  It  seems 
highly  probable  that  S.  avus  is  related  to  the 
generalized  S.  leptorhynchus  based  mainly  on 
meristic  characters.  The  low  trunk  ring  and  high 
tail  ring  counts  of  S.  emeritus  are  unlike  those 
seen  in  living  eastern  Pacific  Syngnathus  spp. 

Syngnathus  auliscus  is  the  only  truly  tropical 
member  of  the  genus  Syngnathus  in  the  eastern 


Pacific.  The  characteristic  spination  of  young  S. 
auliscus  can  also  be  seen  in  young  S.  fuscus 
from  the  northwestern  Atlantic.  This  spination 
has  also  been  reported  (D'Ancona  1933)  for  the 
Mediterannean  S.  agassizi  and  S.  phlegon.  Spi- 
nation in  young  Syngnathus  may  be  indicative 
of  a  distinct  lineage  within  the  genus.  A  system- 
atic study  of  the  juveniles  of  Syngnathus  spp. 
is  definitely  called  for. 

Syngnathus  auliscus  is  not  uncommonly  taken 
in  southern  California.  It  is  apparent  that  this 
species  is  little  affected  by  the  tropical-temper- 
ate transition  between  about  Sebastian  Vizcaino 
Bay  and  Magdalena  Bay.  This  distribution  pat- 
tern is  shown  by  other  fishes  such  as  Paralabrax 
maculatofasciatus  (distribution  in  Miller  and 
Lea  1972).  However,  in  San  Diego  Bay  the 
warm-water  effluent  produced  by  the  San  Diego 
Gas  and  Electric  power  plant  provides  an  ideal 
environment  for  tropical  species.  As  a  result 
there  is  a  large  population  of  5.  auliscus  in  San 
Diego  Bay,  and  the  typical  California  bay 
species  S.  leptorhynchus  is  now  less  abundant. 

The  genus  Bryx  is  a  circumtropical  genus  with 
only  B.  arctus  and  B.  heraldi  having  temperate 
populations.  There  are  eight  species  in  the  New 
World;  one,  B.  balli,  is  in  Hawaii,  B.  banner! 
is  in  the  Indo-West  Pacific,  and  B.  darrosanus 
is  in  the  Indian  Ocean.  The  two  subgenera,  Bryx 
and  Simocampus,  were  present  before  the  clo- 
sure of  the  Panamanian  seaway  in  the  late  Plio- 
cene to  early  Pleistocene  (Woodring  1966),  since 
the  eastern  Pacific  and  western  Atlantic  now 
have  members  of  both  subgenera. 

The  subgenus  Bryx  is  composed  of  four 
species,  two  in  the  Atlantic  and  two  in  the  Pa- 
cific Ocean.  This  subgenus  is  presumably  de- 
rived from  the  subgenus  Simocampus  and  is  dis- 
tinguished from  it  by  the  absence  of  the  anal  fin. 
The  two  species  in  each  ocean  are  more  closely 
related  to  each  other  than  either  is  to  the  species 
in  the  other  ocean.  In  fact  the  radiations  in  the 
two  oceans  have  paralleled  each  other.  The  pri- 
mary difference  between  the  two  species  is  the 
relative  length  of  the  snout. 

Without  a  knowledge  of  the  plesiomorphic 
condition  with  regard  to  meristic  characters,  it 
is  impossible  to  speculate  on  the  relationships 
within  the  subgenus  Simocampus.  However, 
the  Juan  Fernandez  endemic  B.  heraldi  was 
probably  derived  from  B.  coccineus  or  their  im- 
mediate common  ancestor.  This  is  unusual  since 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


221 


the  ichthyofauna  on  the  Chilean  islands  of  Juan 
Fernandez  and  San  Felix  has  a  strong  Indo-Pa- 
cific  component  (McCosker  1970,  1971).  The 
eastern  Pacific  B.  arctus  and  B.  coccineus  are 
more  closely  related  to  each  other  than  either  is 
to  members  of  the  genus  in  other  oceans. 

Doryrhamphus  melanopleura  is  an  Indo-Pa- 
cific  species  that  has  apparently  invaded  the 
eastern  Pacific  by  crossing  the  East  Pacific  Bar- 
rier, since  there  are  no  members  of  this  genus 
in  the  Atlantic,  even  though  suitable  habitat 
would  seem  to  be  available.  The  eastern  Pacific 
populations  of  D.  melanopleura  differ  in  color- 
ation from  their  Indo-West  Pacific  conspecifics. 
The  Islas  Revillagigedo  endemic  D.  paulus  has 
the  same  color  pattern  as  the  eastern  Pacific  D. 
melanopleura  and  was  probably  derived  from 
mainland  populations  of  D.  melanopleura. 

The  genus  Leptonotus  is  a  southern  cold-tem- 
perate group  with  species  in  South  Australia 
(Munro  1958),  New  Zealand,  and  southern 
South  America.  This  genus  is  represented  in 
South  America  by  L.  blainvilleanus.  The  mech- 
anism by  which  cold-temperate  organisms  have 
achieved  a  distribution  spanning  the  South  Pa- 
cific Ocean  is  currently  under  debate.  Rosen 
(1974)  has  argued  that  the  distribution  of  at  least 
the  freshwater  galaxiids  covers  most  of  the  com- 
ponents of  the  original  Gondwanaland  mass. 
Thus,  the  distribution  of  southern  cold-temper- 
ate organisms  reflects  the  break-up  of  Gon- 
dwanaland. However,  the  possibility  of  chance 
dispersal  probably  utilizing  the  eastward-flowing 
South  Pacific  current  seems  to  be  more  likely 
for  most  marine  fishes.  This  has  been  docu- 
mented by  McCosker  (1970,  1971)  for  Paraper- 
cis  and  Muraenichthys.  McCosker  pointed  out 
that  most  of  the  groups  showing  this  particular 
distribution  pattern  are  derived  from  Indo-West 
Pacific  groups  dispersing  southward  to  southern 
Australia  and  New  Zealand,  and  the  subsequent 
movement  eastward  of  cold-adapted  forms 
across  the  cold-temperate  South  Pacific  Ocean 
to  Chile.  Eschmeyer  and  Poss  (1976)  have 
shown  that  with  some  cooling  during  the  Pleis- 
tocene, a  number  of  near-surface  sea  mounts, 
reefs,  and  islands  would  have  been  available  to 
eastward-moving  temperate  biota  in  the  south- 
ern Pacific  Ocean.  Although  they  were  dealing 
with  the  scorpionfish  genus  Maxillicosta,  the 
effects  of  Pleistocene  or  earlier  cooling  would 
also  facilitate  dispersal  of  Leptonotus.  The  abil- 


ity of  marine  fishes  to  cross  oceanic  barriers  has 
been  well  documented  (Rosenblatt  et  al.  1972). 
Pipefishes  (e.g.,  5.  pelagicus)  are  known  to  be 
borne  great  distances  by  the  currents.  It  is  there- 
fore likely  that  the  marine  L.  blainvilleanus  was 
derived  from  an  ancestor  from  the  South  Aus- 
tralia-New Zealand  region  by  chance  dispersal 
of  waifs  to  South  America. 

The  relationships  of  Hippocampus  must  await 
a  revisionary  study  of  the  genus  Hippocampus. 
Our  knowledge  of  the  genus  Hippocampus  is  so 
poor  that  it  is  now  impossible  to  determine  the 
exact  relationship  of  H.  ingens  to  other  mem- 
bers of  the  genus.  Since  the  genus  is  circum- 
tropical,  it  is  at  least  pre-Tethyan  in  origin. 

Hipposyngnathus  has  become  extinct  since 
the  Tertiary.  Because  of  our  almost  complete 
ignorance  of  the  relationships  between  genera  of 
living  syngnathids,  it  is  impossible  to  relate  Hip- 
posyngnathus to  living  genera.  The  reason  or 
reasons  for  the  extinction  of  this  genus  remain 
unknown. 

BRANCHIAL  SKELETON 

Previous  investigators  (Duncker  1910,  1912; 
and  Herald  1959)  have  used  the  position  and  de- 
velopment of  the  male  brood  pouch  as  the  di- 
agnostic character  when  defining  genera  and 
subfamilies.  The  method  of  brood  pouch  closure 
was  used  by  Herald  (1959)  in  constructing  a  phy- 
logeny  of  the  Syngnathidae. 

The  few  osteological  studies  which  have  in- 
cluded the  Syngnathidae  have  generally  been 
based  on  only  one  or  a  few  of  the  genera  (Jun- 
gersen  1910;  Banister  1967).  These  studies  were 
therefore  not  adequately  comparative  and  pro- 
vided little  information  on  intrafamilial  relation- 
ships. 

In  order  better  to  characterize  the  genera  and 
subfamilies  within  the  Syngnathidae,  I  have  be- 
gun a  study  of  the  osteology  of  the  syngnathid 
genera.  This  study  is  also  designed  to  be  a  test 
of  the  hypotheses  of  relationships  presented  by 
previous  authors  who  studied  the  brood  pouch. 
Although  the  present  investigation  is  still  prelim- 
inary, data  can  be  presented  at  this  time. 

Because  the  syngnathids  are  such  highly  spe- 
cialized fishes,  many  aspects  of  their  osteology 
(e.g.,  caudal  skeleton)  are  so  reduced  that  they 
can  provide  little  phylogenetic  information. 
However,  the  branchial  skeleton  does  seem  to 
provide  useful  information.  All  the  genera  ex- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


TABLE  9.     SYNGNATHID  BRANCHIAL  FORMULAE. 


Genus 


Bl 


B2 


B3 


HI 


H2 


El 


E2 


E3 


13       Remarks 


GASTROPHORI: 
Heraldia  (1)* 
Maroubra  (1) 
Choeroichthys  (1) 
Dentirostrum  (1) 
Doryrhamphus  (1) 
Dunckerocampus  (1) 
Syngnathioides  (1) 
Nerophis  (1)** 

UROPHORI: 

Leptonotus  (1) 
Parasyngnathus  (1) 
Ichthyocampus  (1) 
Micrognathus  (1) 
Syngnathus  (7)t 
"Corythoichthys"  (All.)  (1) 
5ry*  (Brw)  (3) 
Bry*  (Simocampus)  (2) 
Pseudophallus  (2) 
Hippocampus  (2) 
PhyUopteryx  (1) 
Corythoichthys  (Pac.)  (2) 
Penetopteryx  (1) 


-IT 


+/f 

+/r 


+  =  present. 
-  =  absent. 

r  =  reduced. 

*  Number  in  parenthesis  represents  number  of  species  examined. 
**  Data  from  Jungersen  (1910). 

t  Only  Syngnathus  aciis  has  reduced  HI. 

1  I2&3  with  tooth  plates. 

2  with  additional  I  element. 

3  I2&3  with  tooth  plates. 

4  I2&3  with  tooth  plates. 


amined  so  far  (Table  9)  have  a  full  complement 
of  ceratobranchials  (i.e.,  five  pairs).  The  re- 
maining elements  in  the  branchial  skeleton  are 
variously  lost  or  reduced  within  the  family  and 
possibly  along  distinct  lineages  within  the  fam- 
ily. The  Urophori  in  particular  show  reduction 
and  loss  of  every  element  except  the  cerato- 
branchials. The  structure  of  the  branchial  skel- 
eton is  constant  within  a  genus.  Although  the 
information  provided  by  the  branchial  skeleton 
is  based  on  characters  undergoing  reduction  and 
not  on  uniquely  derived  characters,  it  is  reason- 
able to  use  these  characters  when  defining  re- 
lationships. It  seems  quite  probable  that  an  ele- 
ment which  is  lost  from  such  a  highly  evolved 
structure  would  not  be  regained. 

The  presence  of  well-developed  first  hypo- 
branchials  and  reduced  second  hypobranchials 
and  epibranchials  clearly  distinguish  the  genus 


Bryx  from  Syngnathus  (Fig.  16).  Bryx  was  in- 
cluded by  Herald  (1959)  as  a  subgenus  of  Syng- 
nathus. 

The  loss  of  all  elements  except  the  first  basi- 
branchial  and  second  hypobranchials  in  the  Pa- 
cific species  usually  referred  to  the  genus  Cor- 
ythoichthys is  clear  evidence  that  the  Atlantic 
and  Pacific  species  should  be  placed  in  separate 
genera  (Fig.  17).  In  fact,  Dawson  (1977)  provi- 
sionally referred  Atlantic  species  of  Corythoich- 
thys to  the  catchall  genus  Syngnathus  and  more 
recently  placed  them  in  a  new  genus,  Cosmo- 
campus  (Dawson  1979).  Herald's  hypothesis 
that  Corythoichthys  is  ancestral  to,  or  on  the 
lineage  leading  to,  Syngnathus  is  clearly  erro- 
neous. However,  Atlantic  species  of  Cosmo- 
campus  could  very  well  be  on  the  lineage  leading 
to  Bryx. 

Herald's  hypothesis  that  Leptonotus  and  Mi- 


FRITZSCHE:  REVISION  OF  EASTERN  PACIFIC  SYNGNATHIDAE 


223 


HI-2 


CI-5 


Bl-2 


EI-3 


11-2 


.Imm 


.Imm 


,1mm  , 


FIGURE  16.  Dorsal  view  of  the  branchial  skeleton  of  several  syngnathids  with  the  epibranchials  and  infrapharyngobranchials 
shown  from  their  ventral  aspect.  (A)  Parasyngnathus  elucens,  CAS  13696.  (B)  Bryx  arctus,  SIO  H52-218.  (C)  Syngnathus  cali- 
forniensis,  SIO  H47-180.  Abbreviations:  B — basibranchial:  H — hypobranchial;  C — ceratobranchial;  E — epibranchial;  I — infra- 
pharynogobranchial. 


crognathus  are  derivable  from  Parasyngnathus 
can  neither  be  supported  nor  refuted  on  the  basis 
of  their  branchial  skeletons.  Parasyngnathus 
has  a  primitive  branchial  skeleton  (Fig.  16).  The 
retention  of  the  upper  pharyngeal  tooth  plates 
in  Micrognathus  is  the  only  major  difference 
between  it  and  Parasyngnathus. 

One  interesting  point  is  the  extreme  reduction 
of  the  branchial  skeleton  in  the  gastrophorine 
genus  Syngnathoides  and  in  the  urophorine  gen- 
era Phyllopteryx  and  Hippocampus  (Fig.  17). 
These  three  genera  are  presently  placed  in  sep- 
arate subfamilies  because  of  differences  in  the 
position  and  development  of  the  brood  pouch. 
However,  in  addition  to  the  reduction  of  bran- 
chial elements,  all  three  genera  have  the  head  at 
an  angle  to  the  main  body  axis  and  a  prehensile 
tail  without  a  caudal  fin.  This  evidence  may  in- 
dicate that  the  three  genera  are  much  more 
closely  related  than  has  previously  been  recog- 
nized. 

The  amount  of  variation  in  the  branchial  skel- 
eton is  much  greater  in  the  Urophori  than  in  the 
Gastrophori.  The  Gastrophorine  genus  Heraldia 
is  the  only  syngnathid  genus  with  three  basi- 
branchials,  and  it  might  well  be  the  most  prim- 
itive of  the  syngnathid  genera. 

The  genus  Nerophis  is  usually  considered  to 
be  the  most  primitive  genus  in  the  family.  How- 
ever, the  species  referred  to  this  genus  have  the 
most  specialized  branchial  skeleton,  in  terms  of 
reduction,  in  the  Gastrophori  (Table  9). 


Imm 


Imm 


B 


\ 


Imm 


Imm 


FIGURE  17.  Branchial  skeletons  of  four  syngnathids.  (A) 
Syngnathoides  biaculeatus,  SIO  60-693.  (B)  Phyllopteryx  fol- 
iatus,  SIO  73-361.  (C)  Cosmocampus  brachycephaltis,  CAS 
24025.  (D)  Corythoichthys  intestinalis,  SIO  73-206. 


224 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  6 


The  evidence  as  is  cited  above  indicates  that 
the  structure  and  development  of  the  branchial 
skeleton  in  the  Syngnathidae  is  of  great  utility 
in  determining  relationships.  The  fact  that  rela- 
tionships as  determined  by  the  branchial  skele- 
ton transcend  recognized  subfamilial  boundaries 
is  a  strong  indication  that  the  within-family  re- 
lationships of  this  group  are  far  from  being 
understood. 

ACKNOWLEDGMENTS 

I  thank  the  following  individuals  for  loan  of 
material  and/or  providing  information  on  speci- 
mens: John  Paxton,  AMS;  Alex  Peden,  BC;  Al- 
wyne  Wheeler,  BMNH;  Keith  Thompson,  BOC; 
William  N.  Eschmeyer,  Tomio  Iwamoto,  and 
Pearl  Sonoda,  CAS;  Fernando  Balbontin, 
EMBCh;  C.  E.  Dawson,  GCRL;  Robert  Behrs- 
tock,  HSU;  Norma  Chirichigno-F.,  IMARPE; 
Robert  J.  Lavenberg,  Camm  Swift,  and  Shelton 
Applegate,  LACM;  Myvanwy  Dick,  MCZ;  M. 
L.  Bauchot,  MNHN;  James  Allen,  SCCWRP; 
Leslie  Knapp,  SOSC;  W.  R.  Taylor  and  Susan 
Karnella,  USNM;  Robert  R.  Miller,  UMMZ;  J. 
Moreland,  NMNZ. 

I  thank  the  following  individuals  for  material 
aid  in  the  completion  of  this  work:  Donald 
Dockins  (deceased),  Joseph  Copp,  and  Jan  Pul- 
sifer  provided  curatorial  aid;  James  Zweifel  pro- 
vided the  Laird-Gompertz  analysis;  John  Butler 
raised  the  pipefish  used  in  the  study  of  growth; 
Myvanwy  Dick  supplied  the  figure  of  Hippo- 
campus ingens  collected  at  San  Francisco;  fel- 
low graduate  students  who  discussed  various 
aspects  of  this  problem  with  me  and  provided 
insights  from  their  own  research.  Dr.  Robert 
Lavenberg  provided  assistance  in  preparing  the 
manuscript  for  publication. 

I  am  particularly  indebted  to  Dr.  Richard  H. 
Rosenblatt  who  made  this  study  possible 
through  his  aid  and  advice,  and  to  Dr.  Carl  L. 
Hubbs  (deceased),  who  generously  provided 
data  collected  on  pipefishes  for  over  30  years 
and  for  his  advice,  support,  and  encouragement. 

And  finally,  I  wish  to  thank  my  wife,  Edith, 
for  her  support  and  encouragement  throughout 
the  course  of  this  research.  Without  her  I  am 
sure  this  work  would  never  have  been  complet- 
ed. 

This  study  is  based  on  a  dissertation  submit- 
ted to  the  University  of  California,  San  Diego, 
in  1976,  in  partial  fulfillment  of  the  requirements 
for  the  Ph.D.  degree. 


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CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  7,  pp.  229-285;  30  figs.,  7  tables  July  2,  1980 


A  REVISION  OF  THE  FISH  GENUS  OGCOCEPHALUS  WITH 

DESCRIPTIONS  OF  NEW  SPECIES  FROM  THE  WESTERN 

ATLANTIC  OCEAN  (OGCOCEPHALIDAE;  LOPHIIFORMES) 

By 
Margaret  G.  Bradbury1 

Department  of  Biological  Sciences,  San  Francisco  State  University, 
San  Francisco,  California  94132 


ABSTRACT.  The  New  World  genus  Ogcocephalus  comprises  twelve  species.  Two  species  are  island  endemics  in 
the  eastern  Pacific,  Ogcocephalus  darwini  Hubbs  in  the  Galapagos  archipelago  and  Ogcocephalus  porrectus 
Garman  off  Cocos  Island,  but  the  remaining  ten  are  western  Atlantic  species.  Five  of  the  Atlantic  species  are 
described  as  new:  Ogcocephalus  pantostictus  and  Ogcocephalus  declivirostris  from  the  northern  and  western 
Gulf  of  Mexico,  Ogcocephalus  rostellum  from  the  Atlantic  coast  of  the  southeastern  United  States,  Ogcocephalus 
corniger  also  from  the  Atlantic  coast  of  the  southeastern  United  States  but  ranging  into  the  eastern  Gulf  of 
Mexico,  and  Ogcocephalus  pumilus  from  the  Caribbean  and  coasts  of  the  Guianas.  Ogcocephalus  parvus  Longley 
and  Hildebrand  has  a  wider  range  than  formerly  known,  from  the  coast  of  the  southeastern  United  States  and 
eastern  Gulf  of  Mexico  through  the  Caribbean  to  the  Atlantic  coast  of  South  America.  Ogcocephalus  vespertilio 
(Linnaeus)  has  a  more  restricted  range  than  formerly  thought,  the  coast  of  Brazil  from  the  mouth  of  the  Amazon 
to  the  mouth  of  the  Rio  de  la  Plata.  Ogcocephalus  notatus  (Cuvier  and  Valenciennes)  also  has  a  southerly 
distribution  in  the  western  Atlantic,  the  coast  of  northern  South  America  from  Colombia  to  northern  Brazil. 
The  variable  species  Ogcocephalus  nasutus  (Cuvier  and  Valenciennes)  appears  to  be  allopatric  with  O,  vespertilio 
(Linnaeus),  which  it  most  resembles;  O.  nasutus  ranges  from  the  mouth  of  the  Amazon  through  the  Caribbean 
to  the  Bahamas  and  southeastern  Florida.  Ogcocephalus  cubifrons  (Richardson)  ranges  from  the  Bahamas  and 
the  coast  of  the  southeastern  United  States  into  the  eastern  Gulf  of  Mexico  to  at  least  Pensacola,  Florida,  and 
Campeche  Banks.  The  name  Ogcocephalus  radiatus  (Mitchill)  is  placed  in  the  synonymy  of  Ogcocephalus  cub- 
ifrons. Illustrated  keys,  photographs,  diagnoses,  and  distribution  maps  are  provided. 

INTRODUCTION  grounds.  Prior  to  this  exploratory  period,  spec- 

Ogcocephalids  from  the  western  Atlantic  imens  of  ogcocephalids  were  uncommon  in  col- 
Ocean  have  become  available  in  large  numbers  lections  except  for  those  captured  inshore  by 
during  the  last  two  or  more  decades,  much  of  swimmers  or  divers,  particularly  off  Florida  and 
the  new  material  a  handsome  dividend  from  the  off  islands  in  the  Caribbean.  Few  in  number  and 
work  of  exploratory  vessels  of  the  U.S.  National  little  studied,  these  specimens  were  difficult  to 
Marine  Fisheries  Service  sampling  new  fishing  assign  names  to  because  variation  in  characters 

then  used  in  diagnoses  was  not  understood. 
Variation  in  the  length  of  the  prominent  rostrum 
was  a  frequent  source  of  error.  The  information 

1  Research  Associate,  Department  of  ichthyology,  Caiifor-      developed  in  this  study  is  that  some  species  may 
nia  Academy  of  Sciences.  be  diagnosed  by  their  long  rostrums  at  all  sizes, 

[229] 


230 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


but  the  rostrum  in  other  species  varies  allo- 
metrically  so  that  juveniles  have  long  rostrums, 
but  adults  have  short  ones. 

Analysis  of  morphometric  data,  including  data 
about  the  rostrum,  reveals  that  the  genus  Ogco- 
cephalus  is  represented  in  the  western  Atlantic 
by  ten  species,  five  of  which  are  described  here 
as  new.  More  recently,  new  material  from  the 
eastern  Pacific  has  also  become  available.  Study 
of  this  material  confirms  that  two  derivative 
species  are  present  in  the  eastern  Pacific  (Gar- 
man  1899;  Hubbs  1958). 

Studies  on  the  remaining  genera  of  Ogcoce- 
phalidae  of  the  Atlantic  Ocean  are  underway 
currently.  Discussions  of  the  zoogeography  of 
the  group  and  intrafamilial  relationships  of  all 
the  Atlantic  species  will  be  published  with  the 
results  of  those  studies. 

ACKNOWLEDGMENTS 

This  study  has  been  pursued  intermittently 
since  the  late  1950's.  During  this  time  I  became 
indebted  to  many  colleagues  to  whom  it  is  a 
pleasure  to  extend  my  grateful  thanks  at  last.  I 
deeply  appreciate  the  encouragement  given  me 
by  the  late  Loren  P.  Woods,  who  first  suggested 
this  problem  and  made  available  the  specimens, 
skeletal  material,  and  photographs  of  ogcoce- 
phalids  in  the  Field  Museum  of  Natural  History; 
he  also  read  an  early  version  of  the  manuscript 
and  devoted  a  great  deal  of  time  to  assisting  me. 
My  dear  friend,  the  late  Marion  Grey,  gave  valu- 
able bibliographic  assistance  as  well  as  the  en- 
thusiastic encouragement  that  so  often  charac- 
terized her  relations  with  her  colleagues.  Pearl 
M.  Sonoda  sent  specimens  on  each  of  a  number 
of  occasions  and  cheerfully  gave  much  other  as- 
sistance. Hymen  Marx  discussed  ideas  with  zest 
and  humor  during  one  memorable  Chicago  sum- 
mer. George  S.  Myers  and  the  late  Rolf  L.  Bolin, 
who  were  my  professors  at  Stanford  University, 
gave  valuable  help;  Prof.  Bolin  gave  skillful  crit- 
icism of  an  early  version  of  this  report.  With 
characteristic  generosity,  Giles  W.  Mead  kindly 
made  possible  a  study  trip  to  museums  in  the 
eastern  United  States  and  gave  support  and  en- 
couragement in  various  ways.  The  late  Carl  L. 
Hubbs  gave  advice  about  taking  counts  and 
measurements  and  graciously  examined  for  me 
a  Linnaean  type-specimen  at  Uppsala.  He  along 
with  W.  I.  Follett  and  Lillian  Dempster  helped 
with  the  coining  of  names. 

Photographs  and  radiographs  were  generously 


provided  by  the  following:  Harvey  R.  Bullis,  Jr., 
Bruce  B.  Collette,  William  N.  Eschmeyer,  Da- 
vid W.  Greenfield,  Ake  Holm,  Robert  J.  Lav- 
enberg,  Garnett  W.  Link,  Jr.,  Robert  N.  Lea, 
Giles  W.  Mead,  and  Loren  P.  Woods.  For  their 
valuable  time,  I  am  grateful  to  the  following  peo- 
ple who  loaned  specimens  or  made  arrange- 
ments for  me  to  work  at  their  institutions:  Wil- 
liam W.  Anderson,  Richard  T.  Barber,  Frederick 
H.  Berry,  James  E.  Bohlke,  the  late  Anton 
Brunn,  Harvey  R.  Bullis,  Jr.,  Daniel  M.  Cohen, 
Bruce  B.  Collette,  C.  E.  Dawson,  Myvanwy  M. 
Dick,  Robert  H.  Gibbs,  Carter  R.  Gilbert,  David 
W.  Greenfield,  Jena  Guibe,  Carl  L.  Hubbs,  Rob- 
ert K.  Johnson,  Robert  J.  Lavenberg,  Giles  W. 
Mead,  George  C.  Miller,  Martin  A.  Moe,  Jr.,  C. 
Richard  Robins,  Richard  H.  Rosenblatt,  Jay  M. 
Savage,  Leonard  P.  Schultz,  Robert  L.  Shipp, 
Victor  G.  Springer,  Camm  C.  Swift,  Ethel wynn 
Trewavas,  Vladimir  Walters,  and  Loren  P. 
Woods.  I  thank  Lillian  J.  Dempster  and  W.  I. 
Follett  for  clarifications  on  nomenclatural  ques- 
tions. 

This  investigation  was  supported  by  Public 
Health  Service  Fellowship  No.  8543  from  the 
Division  of  General  Medical  Sciences,  Public 
Health  Service,  and  by  National  Science  Foun- 
dation grant  GB-4777  from  the  Division  of  Bio- 
logical and  Medical  Sciences.  Support  from 
these  agencies  was  essential  for  the  completion 
of  this  work,  and  I  am  deeply  appreciative. 

SPECIAL  TERMINOLOGY 

The  depressed  anterior  portion  of  the  fish  is 
called  the  disk  and  consists  of  the  cranium, 
opercular  apparatus,  and  the  trunk  back  to  the 
pectoral  axillae.  The  lateral  margins  of  the  disk 
are  formed  by  the  opercular  bones,  which  are 
very  elongate  relative  to  the  cranium.  The  cra- 
nium is  of  ordinary  proportions  relative  to  the 
standard  length,  so  the  large  total  size  of  the 
disk  is  due  primarily  to  the  elongate  opercular 
series,  which  sweeps  backward  from  the  crani- 
um and  envelops  the  trunk  on  either  side  to  form 
the  disk. 

The  (usually)  conspicuous  protuberance  frorn 
the  front  of  the  head  dorsal  to  the  nasal  openings 
is  called  the  rostrum.  Appearing  as  a  median 
horn  in  most  species,  it  varies  in  relative  length 
from  short  to  very  long.  It  may  bear  cirri  at  its 
distal  end,  but  its  function  remains  unknown  at 
this  time.  No  skull  bones  are  involved  in  the 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


231 


OUTSIDE  MARGIN 
OF  LATERAL  ETHMOID 


ESCA 


PREMAXILLARY 
SERIES 


FIGURE  1 .  Lateralis  system  of  Ogcocephalus  cubifrons,  275  mm  SL.  Each  lateral-line  scale  indicated  by  pair  of  opposing 
U-shaped  symbols  representing  flaps  of  skin  associated  with  such  scales.  Canals  indicated  by  dashed  line,  (a)  Lateral  view 
showing  lateral  line  beginning  at  A;  portion  of  supraorbital  series  also  shown,  beginning  at  B.  (b)  Oblique  view  of  head  showing 
cheek  series,  A  to  C,  premaxillary  series,  and  subopercular  series,  C  to  E. 


structure  of  the  rostrum;  it  is  formed  solely  of 
modified  scales. 

Just  beneath  the  rostrum  is  a  cavity  lined  with 
scaleless  skin  into  which  the  angling  lure  or  il- 
licium  may  be  drawn.  This  cavity  is  here  called 
the  illicial  cavity.  The  somewhat  bulbous  struc- 
ture seen  within  the  illicial  cavity  is  the  bait  of 
the  angling  apparatus,  or  esca  (Fig.  \b),  a  mov- 
able glandular  appendage  attached  to  the  distal 
end  of  the  illicium.  The  illicium  is  a  bony  rod, 
quite  short,  but  when  extended,  can  be  seen  to 
be  covered  with  scaleless  skin  like  the  skin 
which  lines  the  illicial  cavity;  the  skin  of  the 
illicium  is  thrown  into  accordionlike  folds  when 
the  illicium  is  retracted. 

In  members  of  the  Ogcocephalidae  the  scales 
are  highly  modified  structures  of  three  main 
types  (Bradbury  1967:  fig.  5).  The  simplest  type 
is  a  pyramid-  or  cone-shaped  scale  called  a  tu- 
bercle, typically  with  a  spine  at  its  apex,  al- 
though there  may  be  several.  The  kind  of  scale 
that  characterizes  Ogcocephalus,  called  a  buck- 
ler, is  larger  and  more  complex;  bucklers  are 
cone-shaped,  too,  but  covered  with  small  spi- 
nules  rather  granular  in  appearance  and  ar- 
ranged in  rows  that  radiate  out  from  the  apex  of 
the  buckler.  These  granular  spinules  fill  the 


spaces  between  bucklers.  The  morphological  re- 
lationship between  tubercles  and  bucklers  is  not 
clear;  the  spinules  of  bucklers  often  resemble 
tubercles,  suggesting  that  the  buckler  may  be  a 
compound  structure  consisting  of  one  enlarged 
tubercle  overlain  with  small  tubercles  arranged 
in  the  radiating  pattern,  but  the  matter  cannot 
be  clarified  until  developmental  studies  of  the 
scales  are  undertaken. 

Finally,  besides  tubercles  and  bucklers,  spe- 
cially shaped  lateral-line  scales  occur.  With  ap- 
propriate preparations  it  can  be  shown  that  lat- 
eral-line scales  are  bowl-  or  dish-shaped 
(Bradbury  1967:  fig.  5),  but  under  ordinary  view- 
ing conditions  their  shapes  are  difficult  to  dis- 
cern. However,  they  are  often  made  conspicu- 
ous because  of  associated  flaps  or  fringes  of 
epidermis;  when  epidermal  decorations  are  ab- 
sent, lateral-line  scales  may  often  be  distin- 
guished because  of  the  large  single  neuromast 
lying  in  the  center  of  the  cup  of  each  scale. 

METHODS 

COUNTS. — Vertebral  counts  were  made  from 
radiographs  and  include  the  hypural  plate. 

Scales  of  the  lateral-line  system  can  generally 
be  seen  in  most  species  of  Ogcocephalidae,  al- 


232 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


POSTERIOR 
SWELLING 
OF    THE 
MANDIBLE 


J 


MEDIAL   RIM 

OF 
SUBOPERCLE 


TERMINAL 
PAIR  OF 
MAJOR  DISK 
BUCKLERS 


SUBOPERCLE 


FIGURE  2.  Ventral  view  of  Ogcocephalus  cubifrons,  275 
mm  SL,  showing  disk  margin,  (a)  Length  of  disk  margin  is 
distance  between  two  points  indicated  by  the  two  arrows,  (b) 
Position  of  subopercle  relative  to  disk  margin. 


though  the  canals  are  difficult  to  discern  as  a 
rule.  Thus  it  may  be  difficult  to  follow  and  count 
lateral-line  scales  when  they  are  small  and 
spaced  far  apart  on  the  surface  of  the  disk  where 
they  tend  to  become  lost  in  the  rough  integu- 
ment, but  elsewhere  on  the  body  it  is  generally 
possible  to  see  the  scales  and  follow  the  lines  of 
the  canals  by  using  adequate  magnification.  In 
this  paper  I  have  used  four  series  of  counts  of 
scales  of  the  lateral-line  system  as  taxonomic 
characters. 

1.  Lateral  line  (Fig.  la).  The  lateral  line  begins 
just  posterior  to  the  eye  at  A.  Care  must  be  tak- 
en not  to  include  scales  of  the  supraorbital  series 
which  begins  at  B.  The  line  of  scales  continues 
back  across  the  dorsal  surface  of  the  disk  and 
suddenly  descends  ventrally  just  posterior  to  the 
gill  pore,  continuing  along  the  lateral  sides  of  the 
tail  and  on  to  the  base  of  the  caudal  fin. 

2.  Subopercular  series  (Fig.  Ib).  This  portion 
of  the  cephalic  lateral-line  system  branches  from 
the  preoperculomandibular  canal  at  C  and  con- 
tinues posteriorly  along  the  ventral  side  of  the 
subopercle  to  E.  A  secondary  branch  leaves  the 
subopercular  series  at  D  and  runs  up  onto  the 
dorsal  side  of  the  disk  for  a  short  distance.  Care 
must  be  taken  to  avoid  including  the  first  scale 


of  this  branch  series  in  the  subopercular  lateral- 
line  scale  count.  The  subopercular  lateral-line 
scale  count  was  always  taken  as  the  count  be- 
tween C  and  D  because  the  posterior  section 
(between  D  and  E)  does  not  vary  from  a  count 
of  3.  Thus,  the  subopercular  lateral-line  scale 
count  in  the  figure  is  6. 

3.  Cheek  series  (Fig.  Ib).  I  have  called  the 
section  of  lateral  line  that  extends  in  a  sigmoid 
curve  from  A  to  C  the  cheek  series,  although  its 
topographic  position  suggests  that  it  consists  of 
lateral-line  scales  from  the  infraorbital  canal  plus 
the  opercular  section  of  the  preoperculomandib- 
ular canal.  The  cheek  series  excludes  scales  of 
the  branch  extending  from  B  towards  the  eye. 

4.  Premaxillary  series  (Fig.  \b).  The  premax- 
illary  series  is  a  short  section  of  infraorbital  ca- 
nal just  above  the  upper  lip. 

MEASUREMENTS. — The  highly  specialized 
body  form  of  these  fishes  requires  some  modi- 
fication of  standard  measuring  procedures  as 
pointed  out  by  Hubbs  (1958).  I  have  followed 
his  procedures  in  large  part,  but  made  adjust- 
ments that  permit  comparative  measurements  of 
members  of  other  genera. 

Measurements  were  made  either  with  sharp- 
pointed  steel  calipers  and  a  steel  rule,  or  with 
dial  calipers,  and  were  recorded  to  the  nearest 
0.1  mm;  measurements  of  approximately  15  mm 
or  less  were  taken  viewed  through  a  low-power 
dissecting  microscope.  All  measurements  in- 
volving the  upper  or  lower  jaws  were  taken  from 
the  anteriormost  median  point  of  the  lips  without 
pressing  the  calipers  into  the  lip  tissue.  All  mea- 
surements involving  bucklers  were  taken  to  the 
base  of  the  buckler  and  not  to  its  apex,  unless 
otherwise  stated. 

Standard  length  is  the  distance  from  the  upper 
lip  to  the  base  of  the  caudal  rays.  The  latter  point 
is  difficult  to  establish  in  large  specimens  in 
which  the  integument  is  thick  and  stiff;  the  cal- 
ipers were  pressed  firmly  back  against  the  swell- 
ing formed  by  the  bases  of  the  rays.  Predorsal 
distance  is  the  distance  from  the  center  of  the 
upper  lip  to  the  base  of  the  first  dorsal  ray; 
preanal  distance  is  the  distance  from  the  center 
of  the  lower  lip  to  the  base  of  the  first  anal  ray. 
The  distance  from  the  jaw  to  the  anus  is  the 
distance  between  the  center  of  the  lower  lip  and 
the  center  of  the  anal  opening. 

The  point  of  articulation  of  the  mandible  with 
the  suspensorium  is  visible  externally  as  a  pro- 
nounced swelling  at  the  posterior  end  of  the 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


233 


OUTLINE   OF 
ILLICIAL 
FOSSA 


FIRST  PAIR 
OF  MAJOR 
BUCKLERS 


APEX    OF  ANGLE 

OF  SUBORBITAL 

BUCKLERS 


FIRST  PAIR  OF 
MAJOR  BUCKLERS 


POINT  OF  GREAT- 
EST CONCAVITY 
OF  THE  CRANIUM 


POSTERIOR   SWELLING 
OF  THE  MANDIBLE 


FIGURE  3.  Views  of  head  of  Ogcocephalus  cubifrons,  275  mm  SL,  showing  specific  locations  of  points  used  in  taking 
measurements  of  head.  Head  depth  is  taken  with  one  needle  of  calipers  set  between  bucklers  of  first  major  pair  (a)  and  (b), 
with  other  needle  on  apex  of  swelling  of  mandible  (b).  Width  of  cranium  is  taken  between  points  of  greatest  concavity  just 
posterior  to  orbits  (b);  also  shown  by  arrows  in  (a).  Suborbital  width  is  least  distance  between  orbit  and  apex  of  angle  formed 
by  suborbital  bucklers  (b). 


mandible;  the  measurement  of  the  disk  margin 
was  taken  with  one  point  of  the  calipers  pressed 
against  the  front  of  this  swelling  and  the  other 
point  pressed  against  the  posterior  end  of  the 
subopercle  (arrows,  Fig.  2a).  Care  must  be  tak- 
en in  determining  the  posterior  end  of  the  sub- 
opercle in  specimens  with  a  thick  stiff  integu- 
ment, but  the  tip  of  the  subopercle  can  be  felt 
with  the  caliper  point  just  posterior  to  the  ter- 
minal pair  of  large  bucklers  at  the  angle  of  the 
disk.  This  measurement  of  the  disk  margin  was 
substituted  for  head  length,  which  proved  to  be 
an  awkward  measurement  in  some  species. 
Trunk  width  is  the  "width  behind  P,  [pectoral] 
fins"  (Hubbs  1958)  which  "is  taken  where  the 
body  margin  reaches  a  maximum  curvature  near 
the  solid  part  of  the  urosome,  behind  the  disk." 

Length  of  rostrum  was  taken  as  the  distance 
from  the  dorsal  rim  of  the  posterior  nostril  to 
the  distal  tip  of  the  terminal  spine  of  the  rostrum. 
The  nostril  landmark  was  chosen,  in  spite  of 
being  off-center,  because  it  is  in  precisely  the 
same  relative  position  in  all  species.  Although 
the  rostrum  appears  to  be  highly  variable  in 
length  in  Ogcocephalus,  results  of  this  study 
show  that  some  of  the  variation  results  from  al- 
lometric  growth  patterns.  When  this  allometry 
is  borne  in  mind,  the  length  of  the  rostrum 
proves  a  useful  distinguishing  feature  in  these 
fishes. 

Width  of  cranium  (Fig.  3«)  corresponds  to 
"skull  prominence  width"  of  Hubbs  (1958).  It 
is  the  distance  taken  between  the  "points  of 


greatest  concavity"  which  lie  just  posterior  to 
the  orbits  (Fig.  3a,  arrows,  and  36).  Head  depth 
is  the  oblique  distance  from  a  median  point  on 
the  roof  of  the  cranium  between  the  first  pair  of 
major  bucklers  (Fig.  3a)  to  the  apex  of  the  cone- 
shaped  swelling  formed  by  the  mandible  where 
it  articulates  with  the  suspensorium  (Fig.  3&). 
The  first  pair  of  major  bucklers  invariably  occurs 
just  posterior  to  a  noticeably  soft  spot  on  the 
dorsal  surface  of  the  cranium,  which  is  actually 
the  fossa  into  which  the  illicium  retracts  (Fig. 
3fl).  Although  off-centered,  this  measurement  of 
head  depth  taken  as  described  is  a  useful,  re- 
peatable  measurement.  The  eye  measurement  is 
the  greatest  diameter  of  the  bony  orbit;  since 
there  are  no  circumorbital  bones,  the  lateral  pro- 
cess of  the  sphenotic  bone  forms  all  there  is  of 
a  posterior  bony  margin  of  the  orbit.  The  rear 
needle  of  the  calipers  was  always  pressed 
against  this  bone.  Suborbital  width  is  the  least 
distance  between  the  margin  of  the  cornea  (well 
marked  by  its  rim  of  tubercles)  and  the  apex  of 
the  sharp  angle  formed  by  the  suborbital  buck- 
lers just  above  the  angle  of  the  lips  (Fig.  3b). 
Lateral  ethmoid  width  is  the  least  distance  be- 
tween the  outside  margins  of  the  lateral  ethmoid 
bones,  which  are  the  bones  that  form  the  sides 
of  the  illicial  cavity  (Fig.  \b).  Mouth  width  is  the 
greatest  overall  width,  i.e.,  the  distance  between 
the  posterior  tips  of  the  maxillaries.  Taking  this 
measurement  and  the  following  one  require 
some  care  because  the  maxillary  tips  are  usually 
concealed  beneath  the  suborbital  integument. 


234 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


The  length  of  the  jaw  is  taken  from  the  median 
point  of  the  upper  lip  to  the  posterior  tip  of  the 
maxillary.  The  snout  length  measurement  is  the 
least  distance  from  the  anterior  bony  margin  of 
the  orbit  to  a  point  on  the  median  line  just  above 
the  upper  lip  and  exactly  at  the  base  of  a  strip 
of  scaleless  skin  lying  between  the  two  armored 
halves  of  the  snout.  Length  of  dorsal  and  length 
of  anal  are  the  lengths  of  these  fins  from  the  base 
of  the  first  ray  to  the  tip  of  the  longest  ray  with 
the  fin  depressed. 

The  patterns  of  dentition  in  species  of  Ogco- 
cephalus  show  little  variation  from  the  pattern 
described  for  O.  darwini  by  Hubbs  (1958).  Brad- 
bury (1967)  described  the  dentition  in  the  genus 
and  compared  it  with  patterns  in  other  genera 
with  the  use  of  alizarin-stained  and  cleared  spec- 
imens. Since  there  are  no  readily  discernible  dis- 
tinctions in  the  dentitions  of  species  of  Ogco- 
cephalus  that  are  taxonomically  useful,  features 
of  dentition  are  omitted  from  this  report. 

As  is  often  the  case  with  endeavors  of  this 
kind,  sampling  was  not  random,  and  specimens 
were  studied  as  they  came  available  regardless 
of  their  size  or  the  locality  from  which  they 
came.  Usually  habitat  differences  were  not 
known.  Meristic  differences  proved  to  be  less 
useful  for  discriminating  among  populations 
than  body  proportions  and  color  patterns,  and 
the  latter  were  used  repeatedly  in  diagnosing 
specimens.  It  was  convenient  to  express  body 
proportions  as  ratios,  and  these  ratios  are  used 
in  the  species  diagnoses.  However,  for  system- 
atic comparisons,  proportions  are  also  given  in 
terms  of  percent  of  standard  length  in  tabular 
format.  Counts  of  pectoral  fin  rays  and  the  var- 
ious lateral-line  scale  series  were  made  on  both 
right  and  left  sides,  and  counts  from  both  sides 
are  included  in  the  tables;  most  of  the  tables 
present  these  data  in  right-and-left  pairs. 

Abbreviations  for  fish  collections  are  as  fol- 
lows: AMNH,  American  Museum  of  Natural 
History;  ANSP,  Academy  of  Natural  Sciences 
of  Philadelphia;  BMNH,  British  Museum  (Nat- 
ural History):  CAS,  California  Academy  of  Sci- 
ences; FMNH,  Field  Museum  of  Natural  His- 
tory; FSBC,  Florida  State  Board  of  Conservation 
Marine  Laboratory;  GCRL,  Gulf  Coast  Re- 
search Laboratory  Museum;  LACM,  Los  An- 
geles County  Museum;  MCZ,  Museum  of  Com- 
parative Zoology,  Harvard  University;  MNHN, 
Museum  National  d'Histoire  Naturelle,  Paris; 
SIO,  Scripps  Institution  of  Oceanography;  SU, 


Stanford  University  (this  collection  is  now 
housed  in  the  California  Academy  of  Sciences 
and  therefore  listed  as  CAS-SU);  UF,  Florida 
State  Museum,  University  of  Florida;  UMML, 
Marine  Laboratory,  University  of  Miami; 
USNM,  U.S.  National  Museum  of  Natural  His- 
tory, Smithsonian  Institution;  ZMUC,  Univer- 
sitetets  Zoologiske  Museum,  Copenhagen. 

Ogcocephalus  Fischer 

Ogcocephalus  FISCHER,  1813:78  [type-species  Lophius  ves- 
pertilio  Linnaeus,  by  subsequent  designation  of  Jordan  and 
Evermann  1896:511];  BRADBURY  1967. 

Malthe  CUVIER,  1816:311  (part)  [type-species  Malthe  vesper- 
tilio  Cuvier  and  Valenciennes  (non  Linnaeus),  1837,  by  sub- 
sequent designation  of  Gill  1878:232]. 

Malthea  (or  Malthaea)  CUVIER  AND  VALENCIENNES,  1837:440 
[emended  spelling  of  Malthe  and  therefore  taking  the  same 
type-species]. 

Oncocephalus  JORDAN,  1895:506  [unjustified  emendation  of 
Ogcocephalus  Fischer]. 

Onchocephalus  GILL,  in  GOODE  AND  BEAN,  1896:498  [unjus- 
tified emendation  of  Ogcocephalus  Fischer]. 

Of  the  two  unjustified  emendations  of  Ogco- 
cephalus which  have  appeared  in  the  literature, 
at  least  one  had  the  objective  of  correcting  an 
alleged  improper  transliteration  from  the  Greek. 
Jordan  and  Evermann  (1898:2736)  state  with  re- 
gard to  the  original  spelling  of  Ogcocephalus, 
"properly  written  Oncocephalus,  but  Fischer 
chose  the  above  monstrous  spelling."  The  sec- 
ond emended  spelling  may  have  been  introduced 
for  the  same  reason,  but  in  fact  no  reason  was 
put  forward.  In  any  event,  the  original  spelling 
of  the  name  is  a  correct  original  spelling,  and 
the  name  stands  as  Ogcocephalus  Fischer  (see 
also  Briggs  1961). 

DIAGNOSIS. — A  genus  of  Ogcocephalidae  with 
gill  rakers  in  the  form  of  oval  pads  which  are 
studded  with  teeth  resembling  the  fine  teeth 
elsewhere  in  the  buccal  cavity  (Bradbury  1967; 
fig.  7).  Gills  2Vi,  none  on  the  first  arch;  complete 
holobranchs  on  the  second  and  third  arches,  a 
hemibranch  on  the  fourth  arch.  Scales  on  ventral 
surface  of  tail  consisting  of  numerous,  tiny, 
closely  spaced  tubercles;  if  any  bucklers  occur, 
they  are  scattered  or  may  occur  in  a  single  short 
median  row,  but  in  any  case,  never  are  arranged 
in  two  longitudinal  rows  that  cover  the  entire 
ventral  surface  as  in  Zalieutes  or  Malthopsis. 
Esca  distinctly  trilobed,  with  a  median  dorsal, 
somewhat  leaf-shaped,  lobe  and  two  ventral 
lobes  arranged  symmetrically  below  the  median 
lobe  (Bradbury  1967:  fig.  3). 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


235 


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DESCRIPTION.— flo Jy  /orm.  Body  depressed 
anteriorly,  forming  a  disk  which  is  subtriangular 
in  outline.  Cranium  well  elevated  above  general 
slope  of  disk,  proportions  between  width  of  cra- 
nium and  head  depth  showing  little  variation 
within  genus  (width  of  cranium  into  head  depth 
1.4-2.1).  Also,  there  is  little  variation  in  length 
of  snout  and  width  of  suborbital  relative  to  width 
of  cranium  (suborbital  width  into  width  of  cra- 
nium 1.0-1.9;  snout  length  into  width  of  cranium 
1.4-2.4).  Rostrum  variable  in  length,  from  a  rel- 
atively small  knob  to  a  long  median  horn.  Gill 
pores  opening  dorsally,  lying  in  the  axillae  a 
short  distance  in  advance  of  rear  margin  of  disk. 
No  membranes  connecting  pectoral  peduncles 
with  trunk  as  occur  in  Halieutichthys.  Ventral 
surface  of  disk  flat. 

Tail  stout  and  muscular,  relatively  wide  at  its 
junction  with  disk,  tapering  to  a  narrow  caudal 
base.  Ventral  surface  of  tail  flattened  so  that  a 
cross  section  through  anterior  half  of  tail  would 
be  nearly  a  half  circle.  Predorsal  distance  1.4- 
1.7  in  standard  length;  preanal  distance  1.2-1.4 
in  standard  length.  Anus  slightly  in  front  of  a 
line  drawn  between  posterior  tips  of  pectoral 
elbows  when  pressed  against  trunk;  distance 
from  jaw  to  anus  1.6-1.9  in  standard  length. 

Head  Features.  Eyes  directed  laterally  and  a 
little  anteriorly  and  downward.  Cornea  much 
smaller  than  orbit,  skin  extending  from  cornea 
around  eyeball  studded  with  small  tubercles. 
Depending  upon  its  condition  at  time  of  pres- 
ervation, iris  shows  a  more  or  less  pronounced 
pupillary  operculum  (except  in  O.  notatus)  on 
its  ventral  margin,  giving  pupil  an  approximate 
kidney  shape.  Sometimes  a  less  noticeable  dor- 
sal operculum  (again  excepting  O.  notatus)  ex- 
tends downward  towards  its  ventral  mate  so  that 
pupil  is  dumbbell-shaped;  presumably  these 
opercula  regulate  pupil  size,  but  I  know  of  no 
experiments  that  demonstrate  this  function. 

Anterior  nostril  round,  the  aperture  encircled 
by  a  rim  of  skin  which  often  forms  a  short  tube; 
posterior  nostril  slitlike,  larger  than  anterior 
nostril.  Jaws  very  protractile,  mouth  moderate 
to  small,  usually  semicircular,  its  width  2.6-5.2 
in  length  of  disk  margin  except  in  O.  notatus, 
which  has  a  relatively  wide  mouth  and  in  which 
the  ratio  is  1.8-2.5.  In  normal  position,  posterior 
ends  of  maxillaries  concealed  beneath  the  sub- 
orbital  bucklers.  Upper  jaw  terminal  or  subter- 
minal  relative  to  snout;  lower  jaw  included;  gape 
oblique. 


236 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


TABLE  2.     FREQUENCY  DISTRIBUTIONS  FOR  DORSAL  AND  ANAL  FIN  RAY  COUNTS  AND  VERTEBRAL  COUNTS  IN  SPECIES  OF 
Ogcocephalus. 


Dorsal  fin 


Anal  fin 


Vertebrae 


1               2 

3 

4 

5 

6              3 

4 

18 

19 

20 

21 

notatus 

1 



35 

17 

2              5 

50 

6 

11 





darwini 

—            — 

1 

16 

1 

2 

16 

— 

14 

3 

— 

porrectus 

—            — 

11 

22 

— 

—            — 

33 

2 

14 

— 

— 

vespertilio 

—            — 

3 

26 

8 

—            — 

37 

3 

15 

— 

— 

nasutus 

1 

1 

57 

11 

4 

66 

3 

30 

— 

— 

cubifrons 

—            — 

3 

23 

10 

1 

35 

— 

5 

14 

1 

pantostictus 

—            — 

3 

23 

11 

—            — 

37 

— 

2 

12 

3 

rostellum 

—            — 

5 

20 

1 

1 

25 

— 

5 

17 

2 

corniger 

—            — 

7 

30 

1 

1 

37 

3 

16 

— 

— 

parvus 

1 

1 

64 

3 

2 

67 

3 

9 

1 

— 

declivirostris 

—            — 

1 

39 

13 

4 

49 

1 

14 

28 

— 

pumilus 

1 

9 

37 

3 

28 

21 

— 

14 

— 

— 

Fins.  Dorsal  fin  small,  placed  far  posteriorly 
on  tail  to  rear  of  level  of  anus;  dorsal  rays  usu- 
ally 4-5,  unbranched.  Anal  fin  "like  a  narrow 
lappet"  (Hubbs  1958)  or  a  prop  for  the  tail,  usu- 
ally fleshy,  particularly  along  anterior  edge  and 
tips  of  first  two  rays.  Apparently  this  fin  lies 
twisted  on  its  side  while  fish  rests  on  the  sub- 
strate. Typically  the  anal  fin  has  4  unbranched 
rays,  but  O.  pumilus  frequently  has  only  3  (Ta- 
ble 2).  Pectorals  sturdy,  variable  in  shape  from 
relatively  long  and  narrow  to  broad  and  fan 
shaped;  pectoral  rays  with  skin  especially  fleshy 
on  distal  ventral  surfaces  in  9  of  the  12  species, 
this  fleshy  tissue  not  developed  on  ventral  sur- 
face of  pectorals  in  very  small  specimens  as  a 
rule.  Pectoral  rays  10-15,  unbranched  (Table  3). 


Pelvics  also  sturdy,  narrow  at  their  base  for 
more  than  half  their  length,  then  abruptly  flared 
distally  with  tips  of  rays  very  fleshy  in  most 
species,  so  much  so  in  adults  of  some  species 
that  fin  resembles  a  small  paddle.  Pelvic  fin 
count  always  1,5;  rays  unbranched.  Caudal  fin 
with  rounded  margin  and  9  rays,  at  least  6  of 
which  are  branched;  formula  either  i,7,i  or  i,6,ii. 
Teeth.  Teeth  conical,  retrorse,  villiform,  in 
bands  on  jaws,  in  broad  plates  on  tongue,  and 
in  smaller  patches  on  prevomer  and  palatines. 
Gill  rakers  in  the  form  of  oval  plates,  variable 
in  size  but  always  bearing  teeth  similar  to  those 
on  jaws  and  in  buccal  cavity.  Patches  of  slightly 
larger  teeth  on  second  and  third  pharyngeobran- 
chials,  these  patches  always  well  developed 


TABLE  3.     FREQUENCY  DISTRIBUTION  OF  PAIRS  OF  PECTORAL  FIN  RAY  COUNTS  IN  SPECIES  OF  Ogcocephalus. 

Paired  pectoral  ray  counts 


18 

tf 

H 

H 

H 

« 

fi         W         tJ         tt         ft             other 

X 

notatus 
darwini 
porrectus 

— 

— 

— 

— 

2 

22           3         18           1         —                ff 
8           6         14 

5         23           2           2 

13.4 
14.6 
14.0 

vespertilio 

— 

— 

— 

— 

1 

4           5         28           4           4 

14.0 

nasutus  (South  Amer.)        — 

— 

i 

— 

3 

4 

20          6           5         — 

13.1 

nasutus  (Cent.  Amer.)        — 

— 

i 

i 

17 

2 

4         

12.1 

nasutus  (W.  Indies) 

— 

— 

i 

24 

4 

9         —           1         —         —                — 

12.3 

cubifrons 

— 

— 

i 

24 

12 

44           9           2         -                        H.tt.tt 

12.7 

pantostictus 

— 

i 

3 

17 

4 

21         —           1         —        — 

12.5 

rostellum 

— 

— 

1 

2 

2 

32           3           3         —        — 

13.0 

corniger                                   2 

5 

25 

2 

3 

— 

—        —        _        _        _                _ 

11.0 

parvus  (northern)                 23 

6 

12 

— 

— 

— 

—        _        —        _        _                _ 

10.4 

parvus  (southern) 

— 

11 

3 

1 

— 

_        —        —        _        _                _ 

11.2 

declivirostris                          2 

4 

41 

2 

2 

— 

A 

11.0 

pumilus 

1 

30 

8 

9 

— 

—       —       —       —       _             — 

11.3 

BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


237 


I  I  I  I  I 


oo  oo  r~   — 


VD    O\    (M    CM 


—  \o  oo  oo 


<N    I/I    m    I/I    •*    \O 


00    <N    —    O\    fN    ON    O\ 


—    00       I      f)    O 


~   r»>   Tt   ao 


g: 

! II  111 I  ill 


even  in  species  in  which  gill  rakers  are  smallest 
(Bradbury  1961). 

Integument.  Most  species  with  dermal  cirri, 
but  these  variable  among  individuals  of  the  same 
species;  also,  cirri  are  probably  fragile  and  most 
likely  do  not  preserve  uniformly  well,  so  their 
variability  is  difficult  to  assess.  In  any  event, 
when  they  appear  in  preserved  materials,  they 
may  occur  singly  or  in  clusters  or  thick  fringes, 
most  often  in  association  with  lateral-line  sys- 
tem, particularly  on  face,  disk  margin,  and  lat- 
eral sides  of  tail.  Scales  consisting  of  close-set 
tubercles  and  bucklers,  their  bases  sometimes 
slightly  overlapping,  forming  a  heavy  armor. 
Large  bucklers  in  dense  groups  over  cranium 
and  face  and  around  disk  margins,  where  they 
may  be  ankylosed  to  underlying  bone.  Heavy 
bucklers  also  on  dorsal  surface  of  trunk  and  tail 
where  they  form  a  median  band  which  varies  in 
width,  extending  in  some  species  down  sides  of 
tail  to  lateral  line.  All  species  with  large  bucklers 
scattered  elsewhere  over  dorsal  surface  of  disk 
and  pectoral  peduncles  with  tubercles  and  small 
bucklers  interspersed  among  them.  Tubercles 
always  invest  skin  covering  surface  of  eyeballs 
between  cornea  and  edges  of  bony  orbits.  Ven- 
tral surface  of  disk  invariably  covered  with  fine 
tubercles  so  that  skin  looks  and  feels  like  sha- 
green. Ventral  surface  of  tail  similarly  covered, 
but  there  may  be  a  short  median  line  or  a  scat- 
tering of  small  bucklers.  Dorsal  and  pectoral  fins 
may  or  may  not  have  minute  tubercles  running 
out  along  rays,  but  the  fleshy  anal  and  pelvic 
fins  always  have  fine  tubercles  covering  at  least 
basal  l/3  or  l/4  of  fin,  except  in  O.  notatus  in 
which  fins  are  nearly  naked.  Caudal  fin  with 
thick  bands  of  small  bucklers  and  tubercles  run- 
ning out  along  rays,  but  in  alternating  fashion  so 
that  rays  1,  3,  5,  etc.,  have  their  thickest  cov- 
ering on  one  side  of  the  fin  while  alternate  rays 
have  their  thickest  covering  on  opposite  side, 
permitting  fin  to  be  tightly  folded  in  spite  of  its 
dense  armor. 

Color  in  preservative.  Most  species  with 
markings  on  "wings"  of  disk  and  in  axillae  in 
the  form  of  clusters  of  spots.  Less  commonly, 
tracts  of  spots  or  reticula  occur  along  lateral 
walls  of  tail,  and  in  Ogcocephalus  porrectus  and 
O.  darwini  the  disk  and  tail  markings  have  co- 
alesced into  a  pair  of  prominent  stripes.  Color 
of  dorsal  and  pectoral  fins  variable  from  species 
to  species,  but  color  of  anal  and  pelvic  fins  gen- 
erally the  same  as  color  of  ventral  surface  of 


238 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


TABLE  5.     FREQUENCY  DISTRIBUTION  OF  PAIRS  OF  CHEEK  LATERAL-LINE  SCALE  COUNTS  IN  SPECIES  OF  Ogcocephalus. 

Paired  cheek  lateral-line  scale  counts 


other 


notatus 

43 



— 

— 

— 

3            37             3             — 

10.0 

darwini 

24 

1              4 

15 

2 

1 

i 

7.9 

porrectus 

19 

—            — 

18 

— 

— 

1 

8.0 

vespertilio 

34 

—            — 

— 

2 

31 

1 

9.0 

nasutus  (South  Amer.) 

38 

—            — 

— 

4 

31 

2              1 

9.1 

nasutus  (Cent.  Amer.) 

20 

—            — 

4 

5 

9 

1                        A 

8.7 

nasutus  (W.  Indies) 

37 

—            — 

8 

6 

22 

1 

8.7 

cubifrons 

50 

3 

31 

9 

6 

l                                   — 

8.2 

pantostictus 

39 

1 

2 

10 

9 

13              3                            A 

9.0 

rostellum 

25 

—           — 

19 

5 

1 

—           —           —             — 

8.1 

corniger 

26 

1 

22 

2 

— 

—           _           —             _ 

8.0 

parvus  (northern) 

35 

1              6 

24 

2 

— 

l,i 

7.9 

parvus  (southern) 

15 

1 

11 

3 

— 

—           —           _             _ 

8.1 

declivirostris 

42 

—           — 

36 

5 

— 

i 

8.1 

pumilus 

32 

—           — 

23 

7 

2 

—           —           —             — 

8.2 

body.  Ventral  surface  of  body  usually  much  pal- 
e:*  than  ground  color  of  dorsal  surface.  Dorsal 
and  pectoral  fins  may  have  spotted  or  reticulate 
pattern,  but  anal  and  pel  vies  seldom  have  any 
discrete  spotting  or  banding.  Caudal  fin  mark- 
ings nearly  constant  throughout  genus,  consist- 
ing of  a  dark  basal  third  about  the  same  intensity 
as  ground  color  of  dorsal  surface  of  body,  a  pale 
middle  third,  and  a  dark  distal  third.  Spots 
sometimes  superimposed  on  this  basic  pattern 
in  a  few  species. 

Color  in  life.  Observations  of  life  colors  in 
these  fishes  suggest  that,  as  a  group,  they 


are  remarkably  colorful  if  one  considers  how 
drab  they  become  in  preservative.  In  most  cases 
the  underparts  are  some  shade  of  red:  reddish 
orange,  or  crimson,  or  even  purplish  red,  with 
fins,  lips,  and  buccal  membranes  likely  to  be  col- 
ored a  shade  of  red  as  well.  One  relatively  deep- 
water  form,  O.  pumilus,  is  pinkish  colored  when 
freshly  caught.  Life  colors  are  known  from  too 
few  species,  however,  to  be  able  to  generalize 
yet  about  life  colors  in  these  fishes. 

DISTRIBUTION. — Ogcocephalus  is  confined  to 
the  West  Indian-Panamanian  fauna,  with  ten 
species  in  the  western  Atlantic  Ocean  and  two 


TABLE  6. 
cephalus. 


FREQUENCY  DISTRIBUTION  OF  PAIRS  OF  SUBOPERCULAR  LATERAL-LINE  SCALE  COUNTS  IN  SPECIES  OF  Ogco- 


Paired  subopercular  lateral-line  scale  counts 


N        J 

f         f 

i 

f 

$ 

I 

f         f 

% 

9                10              10 
TO           TO            11 

other            x 

notatus 

44 







3 

4 

3         8 

19 

3       —      — 

i.U.ft         8.5 

darwini 

21         1 

1 

7 

3 

4 

2 

1         .1 

— 

—      —      — 

I               6.5 

porrectus 

31 

—        2 

19 

8 

1 

1 

—      — 

— 

_     _     — 

6.1 

vespertilio 

40 

—      — 

— 

— 

— 

4 

6        8 

13 

5       —      — 

T%,£,A,&        8.6 

nasutus  (South  Amer.) 

38       — 

—      — 

2 

3 

1 

7 

2         9 

6 

4 

M,i,i          8.0 

nasutus  (Cent.  Amer.) 

22      — 

1 

1 

2 

4 

2 

3         3 

1 

2      —      — 

t.i.t            7.6 

nasutus  (W.  Indies) 

37      — 

1 

4 

6 

10 

7 

2         2 

1 

1         1 

1,1             7.2 

cubifrons 

57      — 

—      — 

— 

4 

17 

14 

11         6 

— 

—      _      _ 

t.U.ft         7.5 

pantostictus 

48 

—      — 

— 

— 

— 

— 

3         8 

22 

7         3         1 

U.ft.tt         9.1 

rostellum 

42 

—        3 

9 

9 

12 

4 

1 

— 

_      _      _ 

*,$,l,4          6.6 

corniger 

33       — 

6 

12 

10 

3 

1 

—      _ 

— 

—      _      _ 

$               6.2 

parvus  (northern) 

35         3 

10       15 

4 

1 

— 

1 

—      — 

— 

_      _      _ 

»               5.4 

parvus  (southern) 

15         2 

8         2 

3 

~  * 

— 

— 

—      — 

— 

_      —      _ 

5.1 

declivirostris 

38      — 

—        2 

8 

7 

13 

4 

2      — 

— 

_      —      _ 

M             6.7 

pumilus 

30         1 

4        8 

13 

4 

— 

— 

—      — 

— 

_      _      _ 

5.8 

BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


239 


TABLE  7.     MORPHOMETRIC  PROPORTIONS  FOR  SPECIES  OF  Ogcocephalus.  Body  measurements  are  given  as  thousandths  of 
standard  length. 


Length 

Jaw  to  anus 

Snout  to  dorsal 

of  disk  margin 

Width  of  urosome 

N 

Range 

X 

N 

Range 

X 

N 

Range 

x         N 

Range 

x 

notatus 

29 

.549_.796 

.585 

22 

.633-.680 

.657 

50 

.365-.43S 

.403       36 

.191-.318 

.268 

darwini 

28 

.547-.S36      . 

703 

28 

.5S9-.668 

.639 

29 

.32S-.456 

.385      28 

.155-.244 

.210 

porrectus 

31 

.457-.716      . 

585 

31 

.615-.699 

.647 

35 

.419-.539 

.450      35 

.189-.  277 

.243 

vespertilio 

37 

.542-.617      . 

586 

37 

.635-.733 

.670 

46 

.414_.  490 

.456      37 

.180-.  282 

.231 

nasutus  (South  Amer.) 

11 

.520-.618      , 

564 

14 

.611-.682 

.643 

40 

.419-.506 

.464      40 

.185-.271 

.236 

nasutus  (Cent.  Amer.) 

14 

.535-.62S      , 

571 

25 

.415-.482 

.446        7 

.207-.241 

.221 

nasutus  (W.  Indies) 

8 

.550-.588      . 

566 

9 

.626-.6S4 

.656 

40 

.415-.508 

.465      26 

.160-.  262 

.228 

pantostictus 

5 

.529-.574      . 

558 

5 

.646-.671 

.657 

48 

.420-.483 

.441       25 

.196-.281 

.247 

cuhifrons 

16 

.543-.621       , 

,577 

16 

.613-.684 

.653 

54 

.396-.4S2 

.438       31 

.169-.  294 

.237 

rostellum 

20 

.528-.608      , 

551 

20 

.624-.6S6 

.651 

71 

.405-.493 

.448       19 

.211-.282 

.249 

corniger 

32 

.545-.641 

591 

33 

.641-.732 

.680 

37 

.388-.481 

.441       33 

.192-.258 

.228 

parvus  (northern) 

18 

.480-.652 

603 

17 

.646-.704 

.674 

40 

.4S2-.524 

.486      29 

.183-.  284 

.241 

parvus  (southern) 

7 

.610-.636 

621 

8 

.644-.691 

.666 

15 

.453-.501 

.479       13 

.199-.  280 

.245 

declivirostris 

17 

.540-.601       . 

564 

16 

.605-.679 

.635 

56 

.390-.447 

.419      29 

.193-.342 

.264 

pumilus 

29 

.523-.58S      . 

,547 

29 

.611-.844 

.635 

49 

.39S-.463 

.429      29 

.204-.  264 

.235 

Head  depth 

Width  of  cranium 

Length  of  rostrum 

Width  of  mouth 

N 

Range 

X 

N 

Range 

X 

N 

Range 

x          N 

Range 

x 

notatus 

49 

.198-.  260 

.223 

53 

.121-.185 

.146 

50 

.050-.  195 

.079      53 

.172-.213 

.1% 

darwini 

29 

.196-.291 

,223 

29 

.103-.  155 

.123 

29 

.088-.  181 

.128       29 

.106-.  137 

.121 

porrectus 

35 

.207-.250 

,224 

35 

.120-.  207 

.137 

32 

.131-.164 

.  149      35 

.127-.  153 

.139 

vespertilio 

35 

.224-.255      . 

239 

46 

.104-.  157 

.140 

46 

.169-.293 

.213       37 

.134-.  165 

.152 

nasutus  (South  Amer.) 

41 

.213-.282 

,234 

41 

.127-.  172 

.142 

39 

.079-.230 

.135      41 

.134-.  177 

.149 

nasutus  (Cent.  Amer.) 

24 

.212-.262 

240 

25 

.110-.  150 

.135 

25 

.068-.258 

.136      24 

.122-.  162 

.144 

nasutus  (W.  Indies) 

34 

.208-.262 

229 

40 

.118-.  176 

.133 

37 

.083-.229 

.132       34 

.120-.  164 

.137 

pantostictus 

36 

.200-.250 

,223 

48 

.119-.  155 

.136 

47 

.067-.  170 

.104      40 

.130-.  168 

.142 

cubifrons 

47 

.206-.258 

.225 

52 

.119-.  177 

.141 

51 

.075-.224 

.111       41 

.097-.  141 

.117 

rostellum 

53 

.185-.234 

,204 

71 

.105-.  165 

.130 

69 

.053-.  123 

.073       52 

.100-.  147 

.118 

corniger 

37 

.223-.290 

.251 

38 

.121-.171 

.136 

36 

.189-.267 

.228       37 

.130-.  183 

.152 

parvus  (northern) 

41 

.238-.301 

.276 

41 

.136-.  180 

.157 

40 

.074-.  152 

.119      41 

.109-.  149 

.123 

parvus  (southern) 

15 

.241  -.270 

,253 

15 

.133-.158 

.146 

15 

.094-.  136 

.118       15 

.095-.  120 

.106 

declivirostris 

56 

.198-.244 

,218 

56 

.107-.  157 

.133 

56 

.029-.  131 

.077      56 

.117-.151 

.133 

pumilus 

49 

.205-.  244 

.222 

49 

.123-.  155 

.137 

49 

.135-.219 

.184      49 

.125-.  158 

.144 

Width 

Depth  of 

Length  of  jaw 

Width  of  eye 

of  interorbital 

caudal  peduncle 

N 

Range 

X 

N 

Range 

X 

N 

Range 

x         N 

Range 

x 

notatus 

52 

.110-.  137 

.124 

53 

.072-.  112 

.086 

53 

.046-.090 

.061       52 

.069-.094 

.082 

darwini 

28 

.081-.  110 

.090 

29 

.081-.  103 

.090 

29 

.050-.080 

.064      28 

.066-.090 

.079 

porrectus 

35 

.087-.  117 

.098 

35 

.078-.  105 

.088 

35 

.055-.090 

.064      34 

.065-.083 

.076 

vespertilio 

46 

.095-.  113 

.106 

46 

.076-.  106 

.095 

46 

.060-.093 

.071       46 

.069-.094 

.080 

nasutus  (South  Amer.) 

41 

.091-.  121 

.101 

40 

.079-.  118 

.091 

41 

.054-.074 

.064      40 

.070-.090 

.081 

nasutus  (Cent.  Amer.) 

25 

.096-.  112 

.104 

25 

.070-.  110 

.094 

25 

.056-.077 

.064        8 

.081-.089 

.085 

nasutus  (W.  Indies) 

40 

.087-.  115 

.098 

40 

.076-.  130 

.088 

40 

.053-.097 

.067      35 

.079-.  122 

.089 

pantostictus 

48 

.085-.  110 

.096 

43 

.068-.  106 

.087 

48 

.054-.080 

.066      47 

.077-.096 

.086 

cuhifrons 

53 

.069-.  100 

.081 

36 

.074-.  116 

.091 

53 

.062-.092 

.075      43 

.078-.  105 

.091 

rostellum 

71 

.070-.  107 

.080 

69 

.073-.  109 

.085 

71 

.044-.083 

.059      59 

.055-.088 

.074 

corniger 

38 

.093-.  125 

.104 

38 

.082-.  124 

.101 

38 

.044-.082 

.064      38 

.061-.087 

.076 

parvus  (northern) 

41 

.082-.  102 

.091 

41 

.098-.  134 

.116 

41 

.042-.076 

.056      40 

.075-.  102 

.083 

parvus  (southern) 

15 

.071-.084 

.078 

15 

.091-.  116 

.103 

15 

.055-.073 

.062       15 

.078-.095 

.087 

declivirostris 

56 

.080-.  108 

.092 

56 

.081-.  107 

.095 

56 

.039-.070 

.050      55 

.077-.  100 

.086 

pumilus 

49 

.070-.  1  1  1 

.093 

49 

.094-.  128 

.106 

49 

.037-.054 

.044      49 

.070-.088 

.079 

240 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


TABLE  7.     Cont. 


Width 
of  suborbital 

Width  between 
lateral  ethmoids 

Length  of  snout 

Length 
of  dorsal  fin 

Length 
of  anal  fin 

N 

Range 

X 

N 

Range 

X 

N 

Range 

X 

N 

Range 

X 

N 

Range 

X 

notatus 

35 

.067-.  112 

.087 

36 

.086-.  110 

.096 

36 

.068-.097 

.080 

48 

.105-.219 

.146 

48 

.114-.  180 

.150 

darwini 

29 

.065-.096 

.084 

29 

.057-.088 

.071 

29 

.059-.083 

.066 

26 

.087-.  121 

.105 

25 

.144-.  191 

.162 

porrectus 

34 

.085-.  103 

.093 

34 

.073-.092 

.080 

34 

.049-.079 

.070 

33 

.087-.  121 

.102 

34 

.150-.  190 

.171 

vespertilio 

46 

.089-.  1  14 

.103 

37 

.064-.084 

.077 

37 

.065-.082 

.074 

34 

.099-.  152 

.123 

35 

.147-.  183 

.167 

nasutus 

(South 

Amer.) 

31 

.084-.  123 

.097 

32 

.069-.  100 

.083 

31 

.063-.084 

.073 

31 

.081-.  147 

.108 

31 

.114-.  178 

.146 

nasutus 

(Cent. 

Amer.) 

8 

.099-.  116 

.104 

21 

.073-.096 

.085 

7 

.076-.  100 

.083 

7 

.095-.  119 

.109 

7 

.140-.  172 

.156 

nasutus 

(W.  Indies) 

40 

.087-.  117 

.103 

34 

.074-.  103 

.083 

34 

.065-.083 

.074 

25 

.089-.  153 

.111 

30 

.123-.  193 

.160 

pantostictus 

31 

.084-.  112 

.097 

25 

.076-.095 

.084 

24 

.065-.084 

.074 

41 

.088-.  160 

.120 

36 

.133-.191 

.164 

cubifrons 

35 

.085-.  110 

.0% 

29 

.072-.096 

.079 

28 

.061-.097 

.073 

41 

.097-.  159 

.126 

29 

.152-.204 

.170 

rostellum 

38 

.070-.097 

.085 

44 

.065-.099 

.076 

20 

.057-.077 

.065 

19 

.088-.  141 

.103 

17 

.122-.  169 

.147 

corniger 

33 

.088-.  112 

.101 

33 

.069-.  1  10 

.081 

33 

.064-.084 

.073 

37 

.072-.  125 

.091 

38 

.124-.  177 

.150 

parvus 

(northern) 

29 

.110-.  136 

.120 

29 

.082-.  117 

.097 

29 

.078-.098 

.089 

36 

.109-.  175 

.145 

39 

.132-.177 

.158 

parvus 

(southern) 

13 

.091-.  110 

.101 

13 

.079-.089 

.084 

13 

.071-.083 

.079 

14 

.120-.  176 

.143 

15 

.143-.202 

.165 

declivirostris 

34 

.069-.  105 

.084 

34 

.068-.  100 

.083 

34 

.058-.084 

.070 

53 

.114-.  186 

.150 

54 

.141-.187 

.166 

pumilus 

29 

.078-.099 

.087 

29 

.069-.094 

.078 

29 

.060-.075 

.067 

43 

.096-.  192 

.132 

44 

.108-.  178 

.150 

in  the  eastern  Pacific.  In  the  Atlantic  the  genus 
occurs  along  the  coast  of  the  United  States  from 
Cape  Hatteras  south  to  the  Gulf  of  Mexico, 
West  Indies,  the  coasts  of  Central  America  and 
South  America  to  Uruguay.  In  the  Pacific  the 
genus  is  known  from  the  vicinity  of  Cocos  Is- 
land, the  Galapagos  Islands,  and  Peru  (but  not 
from  south  of  the  Gulf  of  Panama  as  stated  by 
Lundy  (1956)  or  from  off  Cabo  Corrientes  as 
stated  by  Hubbs  (1958);  of  course,  it  is  possible, 
even  likely,  that  Ogcocephalus  will  be  found 
eventually  at  these  localities,  but  I  know  of  no 
such  records  to  date.  I  suspect  the  error  is  owed 
simply  to  a  misreading  of  coordinates  on  the 
map  of  the  eastern  Pacific  Ocean  concerning  the 
locality  of  the  type-series  of  Ogcocephalus  por- 
rectus (Garman,  1899). 

Relative  to  other  genera  in  the  family,  Ogco- 
cephalus has  the  shallowest  bathymetric  range. 
Although  data  on  depth  distribution  are  still 
scanty  for  some  species,  some  generalizations 
may  be  attempted.  Four  species  appear  to  occur 
most  commonly  at  depths  shallower  than  50  m; 
these  species  are  O.  cubifrons,  O.  pantostictus, 
O.  vespertilio,  and  O.  darwini,  with  cubifrons 
often  recorded  from  depths  of  10  m  or  less.  Two 


species  that  have  their  depth  distribution  cen- 
tered at  50-60  m  are  O.  nasutus  and  O.  notatus, 
and  three  that  have  their  depth  distributions  cen- 
tered at  60-100  m  are  O.  parvus,  O.  pumilus, 
and  O.  declivirostris.  Ogcocephalus  porrectus 
has  been  taken  only  between  100  and  170  m,  but 
the  species  has  been  taken  from  few  localities. 
Finally,  there  are  two  species,  O.  rostellum  and 
O.  corniger,  that  have  relatively  wide  bathy- 
metric ranges  from  30  to  250  m.  Thus,  for  the 
genus  as  a  whole,  most  specimens  have  been 
taken  at  depths  shallower  than  100  m,  but  some 
have  been  taken  as  deep  as  200  m,  and  a  few 
individuals  have  been  taken  from  depths  over 
350m. 


Key  to  Species  of  Ogcocephalus 

Few  species  were  found  to  be  unique  in  the 
characters  studied,  and  constructing  a  key 
proved  difficult.  This  key  will  permit  identifi- 
cation of  most  specimens  of  juvenile  and  adult 
specimens  of  Ogcocephalus,  but  some  couplets 
utilize  characters  that  do  not  discriminate  com- 
pletely between  members  of  the  couplet.  In 
these  cases,  species  intermediate  between  mem- 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


241 


bers  of  a  couplet  are  carried  from  there  through 
both  sides  of  the  key. 

la.  Distribution:  western  Atlantic  Ocean  __   2 
Ib.  Distribution:  eastern  Pacific  Ocean ....    13 

2a.  Pectoral  rays  10/10  to  11/11  or  11/12  (3% 
of  specimens  belonging  to  species  in- 
cluded here  have  pectoral  rays  12/12)  .. 
3 

2b.  Pectoral  rays  12/12  to  15/15  (2%  of  spec- 
imens belonging  to  species  included  here 
have  pectoral  rays  11/12  or  11/11)  6 

3a.  Rostrum  very  long,  length  0.9-1.6  in 
head  depth,  1.6-3.1  in  length  of  disk 
margin 4 

3b.  Rostrum  moderate  to  short,  length  1.7- 
4.0  in  head  depth,  3.2-8.5  in  length  of 
disk  margin  5 

4a.  Anal  rays  3-4.  Body  pale  tan  or  gray 
without  markings  except  dark  spots  oc- 
casionally present  dorsally  on  either  side 
of  disk.  No  fleshy  pads  on  ventral  sur- 
face of  pectoral  rays;  fin  membrane  rel- 
atively thin  and  translucent  (Fig.  4a).  A 
small  species  reaching  70  mm  SL. 
Known  from  northern  Bahamas  through 
Puerto  Rico  and  Lesser  Antilles  to  Sur- 
inam. One  record  from  Honduras 

Ogcocephalus  pumilus 

new  species  (Fig.  26),  p.  272 

4b.  Anal  rays  4.  Dorsal  body  surface  brown- 
ish, evenly  covered  with  small  whitish 
spots  (which  are  the  tips  of  bucklers). 
Distal  ends  of  pectoral  rays  with  thick- 
ened fleshy  pads  on  their  ventral  sur- 
faces; fin  membrane  thick  and  opaque 
(Fig.  4b).  A  moderate-sized  species 
reaching  140  mm  SL.  Known  from  east- 
ern Cuba,  eastern  Gulf  of  Mexico,  and 
eastern  coast  of  United  States  as  far 
north  as  Cape  Lookout,  North  Carolina 

Ogcocephalus  corniger 

new  species  (Fig.  26),  p.  274 

5a.  Rostrum  short,  usually  tilted  upward; 
never  sloping  downward.  Body  surface 
rough,  contours  craggy  with  prominent 
bucklers.  Head  raised  relatively  high 
above  disk,  head  depth  3.1-4.2  in  stan- 
dard length.  Mouth  small,  its  width  2.0- 
2.8  in  head  depth.  Distal  ends  of  pectoral 


FIGURE  4.  (a)  Ventral  view  of  pectoral  fin  of  type  found 
in  O.  notatus,  O.  pumilus,  and  O.  declivirostris.  Skin  rela- 
tively thin,  membranes  translucent,  and  articulations  of  fin 
rays  visible  through  skin,  (b)  Ventral  view  of  pectoral  fin  of 
type  found  in  all  other  species.  Skin  thick,  spongy  along  the 
rays,  especially  distally  where  small  pads  form.  Membranes 
thick  and  elastic.  (These  features  often  undeveloped  in  small 
individuals.) 


rays  with  thickened  fleshy  pads  on  ven- 
tral surfaces;  fin  membrane  thick  and 
opaque  (Fig.  4b).  A  small  species  reach- 
ing 85  mm  SL.  Western  Atlantic  from 
South  Carolina  to  Recife,  Brazil,  except 
absent  from  western  Gulf  of  Mexico  — _ 

Ogcocephalus  parvus 

(Fig.  28),  p.  276 

5b.  Rostrum  horizontal  or  sloping  down- 
ward relative  to  long  axis  of  body.  Buck- 
lers giving  body  a  roughened  surface, 
but  not  craggy  contours.  Head  relatively 
low  to  the  disk,  head  depth  4.3-5.1  in 
standard  length.  Mouth  width  1.4-1.9  in 
head  depth.  Distal  ends  of  pectoral  rays 
without  fleshy  pads;  fin  membrane  thin 
and  translucent  (Fig.  4a).  A  moderate- 
sized  species  reaching  140  mm  SL. 
Known  only  from  northern  and  western 
Gulf  of  Mexico  

Ogcocephalus  declivirostris 

new  species  (Fig.  24),  p.  269 

6a.  Mouth  wide,  its  width  1.8-2.5  in  length 
of  disk  margin,  1.0-1.4  in  head  depth. 
Lateral-line  scales  in  premaxillary  series 
4/4,  in  cheek  series  usually  10/10  (Fig. 
Ib).  Pupil  oval;  no  opercula.  Atlantic 
coast  of  South  America  from  Caribbean 
coast  of  Venezuela  to  mouth  of  Rio  Plata 

in  Uruguay  Ogcocephalus  notatus 

(Fig.  5),  p.  243 

6b.  Mouth  width  2.6  or  more  in  length  of 


242 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


disk  margin,  1.4  or  more  in  head  depth. 
Lateral-line  scales  in  premaxillary  series 
3/3,  in  cheek  series  8/8  to  9/9  or  9/10, 
rarely  10/10  (Fig.  \b).  Pupil  kidney-  or 
dumbbell-shaped  because  of  presence  of 
opercula  (flaps  of  iris  tissue)  encroaching 
over  pupil  7 

7a.  Rostrum  very  long,  length  1.4-3.1  in 
length  of  disk  margin,  0.8-1.7  in  head 
depth  8 

7b.  Rostrum  moderate  to  short,  length  3.2  or 
more  in  length  of  disk  margin,  1.8  or 
more  in  head  depth 10 

8a.  Anal  rays  3-4.  Subopercular  lateral-line 
scales  5/5  to  6/6,  occasionally  6/7.  Inter- 
orbital  relatively  narrow,  2.5-3.7  in 
width  of  cranium.  Distal  pectoral  rays 
without  fleshy  pads  ventrally;  fin  mem- 
brane thin  and  translucent  (Fig.  4a).  A 
small  species  reaching  70  mm  SL.  Known 
from  northern  Bahamas  through  Puerto 
Rico  and  Lesser  Antilles  to  Surinam. 

One  record  from  Honduras 

Ogcocephalus  pumilus 

new  species  (Fig.  26),  p.  272 

8b.  Anal  rays  4,  rarely  3.  Subopercular  lat- 
eral-line scales  7/7  to  9/9  or  9/10,  seldom 
fewer.  Interorbital  wide,  its  width  1.4- 
2.6  in  width  of  cranium.  Distal  ends  of 
pectoral  rays  generally  with  fleshy  pads 
ventrally;  fin  membrane  thick  and 
opaque  (Fig.  46) 9 

9a.  Pectoral  count  most  often  14/14.  Color 
pattern  usually  consisting  of  network  of 
light-colored  lines  on  patches  of  dark 
background,  these  reticulated  patches 
present  dorsally  on  either  side  of  disk, 
sides  of  tail,  bases  of  pectoral  fins,  and 
on  face.  Known  from  mouth  of  Amazon 

River  to  Sao  Paulo,  Brazil 

Ogcocephalus  vespertilio 

(Fig.  5),  p.  247 

9b.  Pectoral  count  most  often  12/12  to  13/13. 
May  be  devoid  of  color  markings,  but 
more  often  with  clusters  of  dark  rounded 
spots,  these  spots  sometimes  with  light 
borders,  present  dorsally  on  either  side 
of  disk  in  some,  extending  on  to  face, 
pectoral  fins,  or  even  entire  dorsal  sur- 
face of  body  in  others  10 


lOa.  Round  dark  spots  occurring  over  entire 
dorsal  surface  of  disk  and  prominent  on 
dorsal  surface  of  pectoral  fins  1 1 

lOb.  Round  dark  spots  either  absent  or,  if 
present,  restricted  to  center  of  disk  on 
either  side;  some  specimens  of  O.  na- 
sutus  from  the  West  Indies  are  darkly 
mottled  and  may  have  networks  of  pale 
lines  on  face,  tail,  and  around  gill  pores. 
Pectorals  without  spotted  pattern,  or  if 
spots  occur,  they  are  dim  and  restricted 
to  the  bases  of  pectorals 12 

1  la.  Mouth  relatively  narrow,  its  width  1.7  or 
more  in  head  depth;  interorbital  wide 
relative  to  the  short  jaw,  interorbital  0.8- 
1.3  in  length  of  jaw.  Subopercular  lat- 
eral-line scales  usually  7/7  to  8/8.  North- 
ern Bahamas  and  coasts  of  southeastern 
United  States  from  Cape  Lookout, 
North  Carolina,  to  Pensacola,  Florida, 
and  south  to  Campeche  Banks,  Mexico 

Ogcocephalus  cubifrons 

(Fig.  18),  p.  258 

lib.  Mouth  wide,  width  1.7  or  less  in  head 
depth;  width  of  interorbital  1.2-1.8  in 
length  of  jaw.  Subopercular  lateral-line 
scales  usually  8/9,  9/9,  or  more.  North- 
ern and  western  Gulf  of  Mexico  from 
Pensacola,  Florida,  to  Tamaulipas, 

Mexico Ogcocephalus  pantostictus 

new  species  (Fig.  23),  p.  264 

12a.  Rostrum  a  well-developed  median  horn. 
Head  often  marked  by  stripes  radiating 
from  eye  in  specimens  from  West  Indies. 
Jaw  relatively  long,  length  1.2-1.6  in 
width  of  cranium.  A  variable  species 
with  a  broad  range  as  understood  here; 
known  from  the  Bahamas,  southeastern 
Florida,  the  Caribbean,  and  northern 
coast  of  South  America  to  mouth  of  Am- 
azon River Ogcocephalus  nasutus 

(Figs.  12,  13),  p.  251 

12b.  Rostrum  extremely  short,  not  produced 
into  a  median  horn  except  in  very  small 
specimens.  No  stripes  radiating  from 
eye,  although  spots  may  occur  on  iris. 
Jaw  short,  length  1.4-1.9  in  width  of  cra- 
nium. Eastern  coast  of  United  States 
from  Cape  Hatteras  to  the  Florida  Keys 

Ogcocephalus  rostellum 

new  species  (Fig.  24),  p.  267 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


243 


13a.  Skin  shagreenlike,  bucklers  not  spiny 
and  protuberant  „  Ogcocephalus  darwini 
(Fig.  29),  p.  279 
13b.  Spiny  bucklers  protuberant;  skin  not 

smooth  and  shagreenlike 

Ogcocephalus  porrectus 

(Fig.  29),  p.  282 

Ogcocephalus  notatus  (Cuvier  and  Valenci- 
ennes) 

(Figure  5) 

[?]  Lophius  histrio  (non  Linnaeus):  ROSENTHAL  1822:pl.  19, 
fig.  2  [skeleton  and  legend;  see  comment  after  synonymy]. 

Malthe  notata  CUVIER,  1829:252  [nomen  nudum]. 

Malthe  angusta  CUVIER,  1829:252  [nomen  dubium;  after  plate 
in  Rosenthal  1822]. 

Malthe  truncata  CUVIER,  1829:252  [nomen  nudum]. 

Malthaea  notata  CUVIER  AND  VALENCIENNES,  1837:453  [Sur- 
inam; holotype  MNHN  4764];  DEKAY  1842:167  [partial  de- 
scription, compiled];  LUTKEN  1866:208  [Maruim,  Brazil; 
description;  truncata  placed  in  synonymy  of  notata]. 

Malthaea  angusta  CUVIER  AND  VALENCIENNES,  1837:454 
[Surinam;  holotype  MNHN  4767]. 

Malthaea  truncata  CUVIER  AND  VALENCIENNES,  1837:454 
[holotype  MNHN  4772]. 

Malthea  notata:  STORER  1846:132  (separate)  [partial  descrip- 
tion, compiled];  CASTELNAU  1855:26,  pi.  12,  fig.  3  [Bahfa. 
Brazil];  GILL  1862:47  [listed,  after  Storer  1846]. 

Malthe  notata:  GILL  1873:14  [range:  southern  Atlantic  coast, 
compiled]. 

Malthe  vespertilio  [?]  (non  Linnaeus):  GILL  1883:556  [cites 
Rosenthal  1822,  in  list  of  osteological  works]. 

Oncocephalus  truncatus:  MIRANDA  Rim  IKO  1915:  on  page 
575  of  unnumbered  text,  2  figs.  [Santos,  Brazil;  description; 
photographs];  1918:154  [synonymy]. 

Ogcocephalus  radiatus:  PUYO  1936:233  [off  Cayenne  River, 
French  Guiana;  description;  figures;  synonymy;  behavior]; 
MENEZES  1964:157  [Brazil;  description;  figures;  compari- 
sons with  Ogcocephalus  vespertilio  (Linnaeus); 
photographs]. 

Ogcocephalus  nasutus  var.  cayennensis  PUYO,  1936:241 
[Cayenne,  French  Guiana;  figures;  holotype  unknown]. 

Oncocephalus  radiatus:  FOWLER  1941:184  [compiled;  after 
Oncocephalus  truncatus  of  Miranda  Ribeiro  1915]. 

Oncocephalus  notatus:  DELSMAN  1941:76  [mouth  of  Amazon 
River;  said  to  resemble  photograph  of  O.  truncata  (Cuvier 
&  Valenciennes)  in  Miranda  Ribeiro  1915]. 

Onchocephalus  radiatus:  PUYO  1949:257  [French  Guiana;  de- 
scription; figures;  synonymy;  behavior]. 

Onchocephalus  nasutus  var.  cayennensis:  PUYO  1949:260 
[Cayenne,  French  Guiana;  description;  figures;  synonymy]. 

Ogcocephalus  truncatus:  LOWE  (McCoNNELL)  1962:679  [Bri- 
tish Guiana;  partial  description;  ecology]. 

Ogcocephalus  nasutus:  CERVIGNON  M.  1966:867  [Venezuela; 
description;  figures;  key];  DAHL  1971:316  [Colombia;  fig- 
ures; key]. 

Ogcocephalus  notatus:  BRADBURY  1967:417  [type-series 
enumerated]. 

The  figure  of  a  skeleton  labeled  Lophius  his- 
trio in  Rosenthal  (1822:pl.  19,  fig.  2)  is  assigned 

to  the  Ogcocephalidae;  the  strongly  depressed 


body  and  the  elongate  pectoral  radials  and  oper- 
cular  bones  are  unmistakably  those  of  an  ogco- 
cephalid.  The  details  of  the  skull,  however,  are 
not  clear  enough  to  identify  the  figure  to  genus. 
The  high  cranium  with  what  appears  to  be  a  sec- 
ondary roof  over  the  illicial  groove  is  character- 
istic of  Ogcocephalus,  and  the  relatively  large 
mouth  suggests  that  the  skeleton  may  be  that  of 
O.  notatus,  but  other  possibilities  cannot  be  ex- 
cluded. I  place  Lophius  histrio  (non  Linnaeus) 
of  Rosenthal,  1822,  in  the  synonymy  of  Ogco- 
cephalus notatus  so  as  to  retain  it  with  the  syn- 
onym Malthe  angusta  Cuvier,  1829,  which  was 
based  on  Rosenthal' s  figure.  Since  the  name  M. 
angusta  Cuvier,  1829,  was  given  in  a  footnote 
with  no  other  designation,  and  since  the  figure 
cannot  be  identified  with  certainty,  the  name 
becomes  a  nomen  dubium.  Two  other  names 
given  in  the  same  footnote,  M.  notata  and  M. 
truncata,  are  without  designations  and  are 
therefore  nomina  nuda. 

All  three  names  were  published  with  desig- 
nations at  a  later  date  (Cuvier  and  Valenciennes 
1837).  Their  holotypes  are  all  representatives  of 
the  same  species,  in  spite  of  the  variation  the 
specimens  exhibit  in  length  of  the  rostrum  and 
number  of  spots  on  the  disk.  Since  M.  notata 
has  page  priority  over  the  other  two  available 
names,  it  is  the  valid  name. 

DIAGNOSIS  AND  COMPARISONS. — Distin- 
guished from  all  other  species  of  Ogcocephalus 
by  its  wide  mouth  (Fig.  6)  and  an  additional  lat- 
eral-line scale  in  the  premaxillary  series  (4  in 
notatus,  3  in  all  other  species).  Four  species  of 
Ogcocephalus  have  geographic  ranges  overlap- 
ping that  of  notatus;  these  are  parvus,  pumilus, 
vespertilio,  and  nasutus.  Of  the  four,  only  na- 
sutus seems  to  occupy  the  same  grounds  and  is 
occasionally  taken  in  the  same  net  hauls  with 
notatus,  but  vespertilio,  which  seems  to  be  an 
inshore  species,  may  also  eventually  be  shown 
to  co-occur  with  notatus.  O.  notatus  is  distin- 
guishable from  nasutus  and  vespertilio  by  the 
shorter  length  of  its  disk  margin  (Fig.  7).  The 
other  two  species,  parvus  and  pumilus,  with 
geographic  ranges  overlapping  that  of  notatus, 
probably  do  not  co-occur  with  notatus  because 
they  have  relatively  deep  bathymetric  distribu- 
tions. O.  notatus  is  distinguishable  from  parvus 
and  pumilus  by  its  higher  number  of  pectoral 
rays  (nearly  always  13  or  more  in  notatus,  12  or 
fewer  in  parvus  and  pumilus,  Table  3). 


244 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


FIGURE  5.  Ogcocephalus  notatus  (Cuvier  and  Valenciennes),  MCZ  45075,  104.5  mm  SL,  from  off  northern  Brazil,  left  top; 
view  efface,  left  bottom,  showing  shelflike  rostrum  (small  specimens  usually  have  short  median  horn).  Ogcocephalus  vespertilio 
(Linnaeus),  CAS-SU  59624,  175.0  mm  SL,  from  off  northern  Brazil,  right  top;  view  of  face,  right  bottom,  showing  reticulated 
pigment  pattern.  O.  vespertilio,  bottom  center,  also  CAS-SU  59624,  150.0  mm  SL,  showing  reticulated  pattern  commonly  found 
on  face  and  iris  in  this  species. 


O.  notatus  is  further  distinguished  from  all 
other  species  in  the  genus  in  having  an  oval- 
shaped  pupil,  whereas  all  the  others  have  the 
iris  modified  by  the  presence  of  opercula  so  that 
the  pupil  is  irregular  in  shape,  approximately 
kidney-  or  dumbbell-shaped.  It  is  inferred  that 
these  opercula  regulate  the  amount  of  light  pass- 
ing through  the  pupil;  if  so,  it  appears  that  O. 
notatus  lacks  this  mechanism.  However,  I  have 
never  observed  this  species  alive.  It  is  possible 


that  appropriate  tests  of  the  iris  in  this  species 
when  subjected  to  bright  illumination  would 
show  that  opercula  are  present  even  though  they 
fail  to  show  in  preserved  materials. 

DESCRIPTION. — Counts  and  measurements 
from  72  specimens,  20.0  to  133.8  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6.  Ogco- 
cephalus notatus  has  relatively  high  meristic 
counts;  the  high  count  for  lateral-line  scales  in 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


245 


nntatus    (5O1          l~ 

. 

darwini   (17) 

r 

•^•-i 

porrectus  (5) 

vespertilio  (37) 

(—  i 

fc-i 

nasutus  (89) 

i  i  1 

pantostictus  (36) 

r 

-^-i 

cubifrons  (53) 

, 

~i 

rostellum  (26) 

r 

—  i 

corniger  (37) 

i—^r 

parvus  (56) 

i  

^-  

• 

declivirostris  (58) 

r 

•     i 

pumi/us  (49) 

a 

h-i 

•.   .   .   . 

Width  of  mouth  into  length  of  disk  margin 

FIGURE  6.  Variation  in  ratio  of  width  of  mouth  into  length  of  disk  margin  in  Ogcocephalus  (horizontal  axis).  Black  bar  = 
2  standard  errors  of  mean,  white  bar  =  2  standard  deviations,  center  vertical  line  =  mean,  and  heavy  horizontal  line  =  range. 
This  figure  was  drafted  before  all  material  of  darwini  and  porrectus  became  available,  but  material  studied  subsequently  does 
not  change  the  point:  O.  notatus  has  a  wider  mouth  than  any  other  species  in  genus. 


cheek  series  (usually  10)  is  diagnostic  for  the 
species  (Table  5).  The  mean  count  of  lateral-line 
scales  in  subopercular  series  in  O.  notatus  (8.5) 
is  exceeded  only  by  pantostictus  (9.1)  and  ves- 
pertilio (8.6)  (Table  6).  The  mean  count  for  pec- 
toral rays  in  notatus  (13.4)  is  less  distinctive, 
resembling  the  nasutus  population  from  off 
South  America  (13.1)  and  rostellum  (13.0),  but 
exceeded  by  darwini,  porrectus,  and  vespertilio 
(all  with  mean  pectoral  counts  of  14.0  or  more) 
(Table  3). 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses. 

Disk  margin  shorter  on  average  than  in  other 
species  of  Ogcocephalus  except  darwini,  its 
length  2.5(2.3-2.8)  in  standard  length.  Tail  wider 
on  average  than  in  other  species,  its  width 
1.6(1.2-2.3)  in  length  of  disk  margin.  Caudal  pe- 
duncle of  average  depth,  2.7(2.4-3.2)  in  head 
depth.  Rostrum  produced  into  a  short  horn  in 
small  specimens,  becoming  extremely  truncated 
and  shelflike  in  larger  specimens;  length  of  ros- 
trum 5.6(2.1-7.9)  in  length  of  disk  margin.  Ap- 
erture of  illicial  cavity  oval  to  subdiamond 
shaped,  wider  than  high  in  specimens  of  medium 
to  large  size,  higher  than  wide  in  specimens 
about  50  mm  SL  or  less.  Cranium  less  sharply 
demarcated  from  disk  than  in  other  members  of 
Ogcocephalus,  its  elevation  above  disk  surface 
(when  viewed  from  front)  presenting  a  convex 
and  gradual  contour,  not  concave  or  abrupt. 


Head  depth  1.8(1.6-2.1)  in  length  of  disk  margin. 
Cranium  broad,  its  width  2.8(2.2-3.3)  in  length 
of  disk  margin.  Eye  1.7(1.4-2.0)  in  width  of  cra- 
nium, prefrontal  width  1.5(1.3-1.7)  in  width  of 
cranium.  Interorbital  space  concave,  moderate- 
ly wide,  its  width  2.4(2.1-2.9)  in  width  of  cra- 
nium, 3.7(2.8^4.6)  in  head  depth.  Mouth  wider 
than  in  other  species  of  Ogcocephalus,  1.1(1.0- 
1.4)  in  head  depth,  2.1(1.8-2.5)  in  length  of  disk 
margin;  length  of  jaw  1.8(1.6-2.2)  in  head  depth. 
Upper  lip  thin,  lower  lip  a  little  fleshy  laterally. 
Dorsal  fin  relatively  long,  its  length  2.8(2.0-3.8) 
in  length  of  disk  margin;  anal  fin  2.7(2.3-3.5)  in 
length  of  disk  margin. 

Integument.  Dermal  cirri  conspicuous  along 
lateral  line  efface,  disk  margin,  and  tail.  Large 
bucklers  present  on  disk  margin  and  median  dor- 
sal area  of  body.  Pectoral  fins  with  tiny  tubercles 
running  out  along  rays  dorsally  but  bare  ven- 
trally.  Dorsal,  anal,  and  pelvic  fins  devoid  of 
tubercles  except  for  a  few  tiny  scattered  ones  at 
bases  of  pelvics  and  anal.  Caudal  fin  as  in  ge- 
neric description.  Skin  covering  the  eyeballs 
bearing  tiny  tubercles;  edge  of  cornea  not  bead- 
ed by  a  row  of  small  bucklers  as  occurs  in  other 
species.  Pectoral  membranes  moderately  thick, 
opaque,  but  no  fleshy  pads  on  ventral  surfaces 
of  ray  tips. 

Color  in  preservative.  Dorsal  surface  uniform- 
ly gray  with  apices  of  bucklers  paler  than  ground 
color.  Distinct  round  spots  with  pale  narrow 
margins  in  a  cluster  on  either  side  of  disk,  1  to 


246 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


o 

f- 

Eg 

a 

d  O 

.E  10 
o> 

o 

£ 
^? 

10 

-o 

°8 

£ 

~& 

c 
o> 

_JO 

—  '  <M 

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£ 

80° 

>  o  o 
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Or 

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.%>•• 

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o  -  vespertilio 

o-  South  Americc 
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40  50  60  70  80  90  100  110 

Standard  length  in  mm. 


120 


130 


140 


FIGURE  7.  Comparisons  between  Ogcocephalus  nasutus  (only  specimens  from  coast  of  South  America),  O.  vespertilio,  and 
O.  notatus  of  relationship  between  length  of  disk  margin  and  standard  length.  O.  notatus  over  75  mm  SL  shown  to  have 
relatively  short  disk  margin. 


7  on  each  side,  occasionally  none,  particularly 
in  very  small  specimens.  One  specimen  41.5  mm 
SL  with  dark  rings  scattered  on  dorsal  surface 
of  body  and  along  sides  of  tail  into  the  axillae. 
Ventral  surface  uniformly  creamy  pale,  chin 
dusky  in  a  few  specimens.  Lips  the  same  pale 
ground  color  as  the  ventral  body  surface.  Iris 
golden  or  dark  gray,  never  with  spots  or  radiat- 
ing lines.  Pectorals  dusky,  shading  to  darker 
gray  distally,  almost  to  black  in  some  very  small 
specimens,  but  tips  of  rays  pale.  Pelvic  and  anal 
fins  the  same  creamy  pale  ground  color  as  ven- 
tral body  surface;  anal  sometimes  with  a  dusky 
spot  distally,  especially  in  small  specimens. 
Dorsal  dusky,  often  with  indistinct  dark  splotch- 
es; an  oblique  black  stripe  slashed  across  dorsal 
fin  rays  and  membrane  in  specimens  about  25.0 
mm  SL  or  less.  Caudal  fin  as  in  generic  descrip- 
tion. 

Color  in  life.  Unknown. 

DISTRIBUTION  (Fig.  8). — Specimens  studied 
ranged  from  Trinidad  on  the  northeastern  coast 
of  South  America  to  the  mouth  of  the  Amazon 
River,  but  the  species  is  known  from  farther 
south  and  also  from  farther  west  into  the  Carib- 
bean. Miranda  Ribeiro  (1915)  reports  one  spec- 
imen, about  125  mm  SL  (his  Oncocephalus  trun- 
catus),  from  Santos,  Brazil;  Cervigon  (1966) 


reports  specimens  from  the  Caribbean  coast  of 
Venezuela  at  Laguna  de  Unare,  and  Menezes 
(1964)  reports  on  a  collection  (his  Ogcocephalus 
radiatus)  from  Aracajii.  Bathymetric  range:  18- 
64  m.  Puyo  (1949)  reports  specimens  from  es- 
tuaries, particularly  of  the  Cayenne  River, 
French  Guiana,  at  depths  of  less  than  2.5  m.  He 
also  reports  one  example  from  a  swampy  up- 
stream section  of  a  creek  8  km  inland  from  Cay- 
enne at  a  depth  of  approximately  1.8  m. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  Data  for  OREGON  stations  (=O)  in  Bullis 
and  Thompson  (1965). 

Trinidad:  ANSP  53381  (1),  Oropuche;  CAS  23928  (2),  O- 
2208. 

Venezuela:  CAS  23929  (2),  O-2223;  CAS-SU  62120  (1),  O- 
2215;  FMNH  65975  (13),  O-2212;  FMNH  65976  (2),  O-2213; 
FMNH  65977  (1),  O-2224. 

Guyana:  ANSP  103629(5),  O-2226;  BMNH  1950.5.15.56(2), 
Georgetown;  BMNH  55.8.28.70  (1),  Demerara;  FMNH  65978 
(24),  O-2225;  FMNH  65979  (77),  O-2226;  FMNH  65980  (1),  O- 
2234;  USNM  197108  (4),  "Cape  St.  Mary." 

Surinam:  CAS-SU  62126  (3),  O-2279,  FMNH  65974  (3),  O- 
2208;  MCZ  45077  (4),  COQUETTE  sta.  193,  6°24'N,  55°03'W, 
24  m;  MNHN  A.4764  (holotype);  MNHN  A.4767  (holotype  of 
Malthaea  angusta  Cuvier  and  Valenciennes);  MNHN  A. 4768 
(1);  MNHN  5259  (1);  USNM  188772  (1),  COQUETTE  sta.  142, 
6°23.5'N,  54°56'W,  26  m;  USNM  188791  (1),  COQUETTE  sta. 
167,  6°18.5'N,  55°28'W,  18  m;  USNM  188781  (8),  COQUETTE 
sta.  2,  6°23'N,  55°05.5'W,  27  m. 

French  Guiana:  USNM  188773  (2),  O-2038. 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


247 


Brazil:  Amapd:  CAS-SU  51215  (5),  O-2058;  FMNH  65971 
(1),  0-2056;  FMNH  65972  (4),  O-2057;  MCZ  45078  (4),  O- 
2059.  Para:  FMNH  65973  (13),  O-2075;  USNM  188797  (1),  O- 
2076. 

Specimens  without  locality  data:  CAS-SU  59629  (1),  Brazil; 
MNHN  A. 4772  (holotype  of  Malthaea  truncata  Cuvier  and 
Valenciennes);  MNHN  5242  (said  to  be  from  New  York,  but 
this  undoubtedly  an  error). 

Ogcocephalus  vespertilio  (Linnaeus) 

(Figure  5) 

Lophius  vespertilio  LINNAEUS,  1758:236  ["American  Ocean"; 
holotype  in  Zoologiska  Institutionen  Uppsala];  HOLM 
1957: 12  [holotype  listed]. 

Malthaea  longirostris  CUVIER  AND  VALENCIENNES,  1837:450, 
pi.  365  [Bahia,  Brazil;  holotype  MNHN  6369];  LUTKEN 
1866:208  [Maruim,  Brazil;  description]. 

Malthea  guacucuja:  CASTELNAU  1855:26  [Bahia,  Brazil;  cited 
as  being  the  same  as  Malthaea  longirostris  Cuvier  and 
Valenciennes]. 

Malthe  vespertilio:  GUNTHER  1861:200  (part)  [partial  descrip- 
tion; places  the  following  names  of  Cuvier  and  Valen- 
ciennes, 1837,  in  the  synonymy  of  vespertilio:  longirostris, 
nasuta,  notata,  angusta,  and  questionably  truncata};  JOR- 
DAN 1885:927  (part)  [longirostris  referred  to  synonymy  of 
vespertilio];  LONNBERG  1896:39  [holotype  listed]. 

Oncocephalus  longirostris:  MIRANDA  RIBEIRO  1915:  on  page 
574  of  unnumbered  text  [Recife,  Brazil;  description]; 
1918:153  [synonymy];  FOWLER  1941:184  [fishes  of  the  coast 
of  Brazil,  compiled]. 

Ogcocephalus  vespertilio:  DE  SOTO  1922:52  [Amazon  R., 
1300-1450  km  upstream];  [?]  CARVALHO  1943:71  [Sao  Pau- 
lo, Brazil;  partial  description];  MENEZES  1964:155  [Brazil; 
synonymy;  description;  photographs];  BRADBURY  1967:417 
[listed];  DE  SCARABINO  1974:306  [mouth  of  Rio  de  la  Plata; 
description;  photographs]. 

DIAGNOSIS  AND  COMPARISONS. — Ogcocepha- 
lus vespertilio  is  distinguished  from  the  majority 
of  species  of  Ogcocephalus  by  its  extremely 
long  rostrum.  Only  two  other  species  in  the  ge- 
nus have  rostrums  that  reach  the  relatively  great 
lengths  seen  in  vespertilio ;  these  two  long-nosed 
species  are  corniger  and  pumilus  (Fig.  9  and 
Table  7),  but  they  are  easily  separated  from  ves- 
pertilio on  the  basis  of  fewer  pectoral  rays  (10- 
12  pectoral  rays  in  corniger  and  pumilus  com- 
pared to  13-15  in  vespertilio).  O.  vespertilio  also 
differs  from  corniger  in  color  pattern,  having 
dark  spots  separated  by  pale  lines  that  usually 
form  a  reticulum  occurring  in  patches  on  the 
face  and  sides  of  the  tail  and  in  the  pectoral  ax- 
illae, while  corniger  is  evenly  covered  by 
closely  set  pale  spots  on  a  darker  background. 
O.  pumilus  is  a  far  smaller  species  than  vesper- 
tilio, reaching  only  about  70  mm  SL;  from  deep- 
er waters  than  vespertilio  (generally  from  waters 
40  m  or  more  in  depth  rather  than  the  inshore, 
shallower  depths  attributed  to  vespertilio),  pu- 


•  -  vespertilio 
o  -  notatus 


FIGURE  8.  Distribution  of  study  material  of  Ogcocephalus 
notatus  and  O.  vespertilio.  O.  notatus  also  known  from  Ca- 
ribbean and  from  Brazil,  and  O.  vespertilio  from  mouth  of  Rio 
de  la  Plata  (see  text). 


milus  is  a  pale  gray  species  generally  lacking 
markings  of  any  kind;  at  the  most,  it  has  only 
light  gray  spots. 

Ogcocephalus  nasutus,  cubifrons,  and  pan- 
tostictus  all  closely  resemble  vespertilio  in  their 
large  size  and  body  shape,  but  the  relative  length 
of  the  rostrum  in  nasutus,  while  highly  variable, 
is  seldom  as  great  as  in  vespertilio  (Fig.  10); 
cubifrons  and  pantostictus  show  allometric 
growth  of  the  rostrum,  with  the  smallest  speci- 
mens having  long  rostrums  and  juvenile  and 
adult  specimens  having  progressively  shorter 
ones.  O.  vespertilio  is  distinguished  from  na- 
sutus by  its  relatively  wide  interorbital  distance 
compared  to  the  distance  between  the  lateral 
ethmoids  (Fig.  1 1)  and  by  the  modal  number  of 
pectoral  rays  (14  in  vespertilio,  12-13  in  nasu- 
tus, Table  3). 

None  of  the  long-nosed  forms  of  Ogcoceph- 
alus are  sympatric  with  vespertilio,  including 
cubifrons  and  pantostictus,  which  are  long- 
nosed  only  when  small.  These  latter  as  well  as 
corniger  are  at  the  northern  edge  of  the  range 


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20  30  4.0 


50 


•JBMMMMMMMaBMMMBBMB  B  MM 


pantost/ctus 


70 


FIGURE  9.  Histogram  showing  ratio  of  length  of  rostrum  into  length  of  disk  margin  (horizontal  scale).  One  square  =  one 
specimen.  White  squares  =  specimens  less  than  50  mm  SL  (not  shown  for  long-nosed  species  vespertilio,  corniger,  and  pumilux, 
nor  small  species  parvus).  This  figure  was  drafted  before  all  material  of  darwini  and  porrectus  became  available,  but  new 
material  does  not  change  general  picture.  (With  inclusion  of  new  material,  data  for  30  darwini  are:  range  2.8  to  3.6,  x  =3.1; 
for  35  porrectus:  range  2.8  to  3.4,  x  =  3.0.) 


for  the  genus  (east  coast  of  the  United  States 
and  the  Gulf  of  Mexico),  while  vespertilio  oc- 
cupies the  southernmost  range  for  the  genus 
(coasts  of  Brazil  and  Uruguay).  O.  pumilus  and 
nasutus  have  intermediate  ranges  (together 
ranging  from  Cuba  and  the  Bahamas  to  Suri- 
nam). Only  nasutus  has  a  range  which,  as  pres- 
ently known,  reaches  almost  to  the  northern 
edge  of  the  range  of  vespertilio,  that  is,  the  vi- 
cinity of  the  mouth  of  the  Amazon  River. 

The  one  species  of  Ogcocephalus  which  is 
sympatric  with  vespertilio  is  the  short-nosed 
form,  notatus,  also  distinguished  from  vesper- 


tilio by  its  wide  mouth  (Fig.  6)  and  higher  num- 
ber of  lateral-line  organs  in  the  cheek  series  (4 
in  notatus  but  3  in  vespertilio). 

DESCRIPTION. — Counts  and  measurements 
from  46  specimens,  52.0-222.5  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6.  Menezes 
(1964)  recorded  the  pectoral  count  for  O.  ves- 
pertilio as  13-14,  but  I  find  a  wider  range  of  12- 
15  with  a  strong  mode  of  14,  which  places  the 
species  second  only  to  O.  darwini  in  high  num- 
ber of  pectoral  rays.  O.  vespertilio  also  has  rel- 
atively high  counts  for  all  lateral-line  scale  se- 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


249 


40 


e 

^ 
*_ 

to 
o 


c 

OJ 


30 


20 


10 


A  =  rostellum 
•  =  pantost/ctus 
a  =  cubifrons 
o  =  nasutus 
®  =  vespertilio 


® 


O 


<D 


O 


«® 


•t 


® 


® 


20  30  40  50 

Length  of  disk  margin  in  mm. 


60 


70 


80 


FIGURE  10.  Comparisons  among  five  closely  similar  species  of  Ogcocephalus  to  show  relationship  of  length  of  rostrum  to 
length  of  disk  margin  (see  also  Fig.  9).  O.  vespertilio  shows  long  rostrum  at  all  stages  of  growth,  cubifrons  and  pantostictus 
show  shortening  of  rostrum  with  growth,  rostellum  shows  short  rostrum  at  all  stages  of  growth.  Length  of  rostrum  in  nasutus 
extremely  variable. 


ries:  subopercular  lateral-line  scale  count  modally 
9  as  in  notatus  and  pantostictus',  modal  count 
for  the  cheek  series  9.  Scales  in  lateral  line  rel- 
atively numerous  as  in  three  other  large  species: 
nasutus,  pantostictus,  and  cubifrons. 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses. Length  of  disk  margin  intermediate 
within  the  range  of  variation  for  the  genus, 
2.2(2.0-2.4)  in  SL.  Width  of  tail  also  interme- 
diate, 2.0(1.5-2.5)  in  length  of  disk  margin;  cau- 
dal peduncle  moderately  deep,  3.0(2.5-3.5)  in 
head  depth.  Rostrum  with  a  thick  base,  propor- 
tionately longer  than  in  any  other  species  save 
corniger,  its  length  2.1(1.5-2.8)  in  length  of  disk 
margin.  Aperture  of  illicial  cavity  elongate,  tri- 
angular, always  higher  than  wide.  Cranium  ris- 
ing steeply  above  the  disk,  becoming  massive  in 
large  adults;  head  depth  1.9(1.7-2.0)  in  length  of 
disk  margin.  Width  of  cranium  3.3(2.9-4.0)  in 
length  of  disk  margin.  The  following  in  width  of 
cranium:  eye  1.5(1.3-1.8),  lateral  ethmoid  width 
1.9(1.6-2.3).  Interorbital  moderate  to  wide,  flat 


posteriorly  but  distinctly  convex  anteriorly, 
grading  into  the  roundness  of  the  rostrum;  in- 
terorbital  width  2.0(1.5-2.4)  in  width  of  cranium, 
3.4(2.6-4.0)  in  head  depth.  Mouth  moderate,  its 
width  1.6(1.5-1.7)  in  head  depth,  3.0(2.7-3.6)  in 
length  of  disk  margin;  jaw  2.2(2.0-2.4)  in  head 
depth.  Upper  lip  fleshy,  of  an  even  width  for  its 
entire  length;  lower  lip  also  fleshy,  thickened 
medially.  Dorsal  fin  of  moderate  size,  3.8(3.0- 
4.6)  in  length  of  disk  margin;  anal  fin  2.7(2.4- 
3.1)  in  length  of  disk  margin. 

Integument.  Dermal  cirri  present  on  tip  of  the 
rostrum,  disk  margins,  and  lateral  sides  of  tail, 
usually  sparse  except  on  rostrum.  Large  buck- 
lers prominent,  arranged  as  in  generic  descrip- 
tion. All  fins  with  tiny  tubercles,  those  on  pec- 
torals restricted  to  the  base,  those  on  ventrals 
and  anal  running  out  from  base  for  about  half 
length  of  fin.  Tubercles  on  dorsal  fin  sparse  ex- 
cept for  anterior  edge  of  fin  where  they  thickly 
encrust  the  first  ray  and  run  out  nearly  to  end 
of  ray.  Caudal  as  in  generic  description.  Skin 
covering  eyeballs  studded  with  tiny  tubercles, 


250 


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20 

e15 

e 

c 

0 

• 

<5> 

interorbital 

0 

0      o     • 

OQ    o 

OQ^)     ^  % 

o  °%CT?  •* 

•fc 

.c 
TJ 

oo   ' 
oo  • 
o       • 

£•' 

o  -  vespertilio 
•  -  nasutus 

o 

f 

5  10  15 

Lateral  ethmoid  width  in  mm., 


20 


FIGURE  11.     Comparison  between  Ogcocephalus  vespertilio  and  O.  nasutus  (only  specimens  from  South  America)  to  show 
relatively  wide  interorbital  distance  compared  to  lateral  ethmoid  width  in  vespertilio. 


the  rim  around  cornea  with  small  bucklers,  but 
these  not  so  prominent  as  in  other  species.  Pec- 
toral membranes  thick,  opaque,  ventral  surface 
of  ray  tips  with  well-developed  fleshy  pads. 

Color  in  preservative.  Ground  color  medium 
brown.  Markings  consist  of  dark  spots  separated 
by  pale  lines  that  form  a  reticulum.  These  mark- 
ings occur  on  face,  often  within  illicial  cavity, 
and  on  esca,  and  generally  extend  in  a  band 
across  each  eye.  On  the  eye,  including  the  iris, 
light  lines  radiate  from  pupil.  The  markings  ex- 
tend posteriorly  from  each  eye  and  are  often 
continuous  on  each  side  with  a  patch  of  mark- 
ings on  each  shoulder.  Similar  markings  extend 
from  axillae  along  lateral  sides  of  tail.  Some- 
times patch  in  axilla  also  continuous  with  shoul- 
der patch  on  each  side.  Ventral  surface  of  body 
uniformly  light  yellowish  brown,  or  sometimes 
chin  and  tail  slightly  darker.  Lips  pale;  rostrum 
with  a  diffuse  dark  subterminal  ring.  Iris  marked 
with  an  extension  of  the  same  pattern  that  cov- 
ers face.  Pectorals  with  reticulate  markings  ba- 
sally,  grading  into  a  dark  border;  rays  tips  pale. 
Dorsal  also  with  reticulate  pattern.  Ventrals  and 
anal  the  same  ground  color  as  ventral  surface  of 
body;  tips  of  ventrals  dusky,  tip  of  anal  dusky 


to  black.  Caudal  as  in  generic  description,  but 
with  reticulate  markings  superimposed  on  basal 
two  thirds. 

Color  in  life.  Unknown. 

DISTRIBUTION  (Fig.  8). — Coasts  of  Brazil  and 
Uruguay  from  Amazon  River  south  to  La  Plata 
River.  Little  bathymetric  data  available;  appears 
to  be  an  inshore  species.  One  specimen  identi- 
fied as  Ogcocephalus  vespertilio  recorded  from 
1300-1450  km  upstream  in  Amazon  River  (de 
Soto  1922). 

MATERIAL  EXAMINED. — The  holotype,  Linnesamling  no. 
163,  Museum  Alstromer-Linne  no.  40,  Uppsala  University, 
examined  for  me  by  C.  L.  Hubbs;  exact  locality  unknown.  All 
other  material  from  Brazil;  numbers  in  parentheses  are  num- 
bers of  specimens.  Para:  USNM  12575  (1),  Amazon.  Mara- 
nhao:  CAS-SU  59627  (1)  and  CAS-SU  59628  (4),  Rio  Bacanga, 
Madre  Deus;  CAS-SU  59630-59632  (3),  Madre  Deus.  Ceard: 
CAS-SU  59616-59620  (5)  and  CAS-SU  59622-59626  (10),  Mu- 
curipe,  Fortaleza;  CAS-SU  59621  (2),  Fortaleza.  Pernambu- 
co:  USNM  104286  (1),  Recife.  Bahia,  all  from  Salvador:  CAS- 
SU  59633  (1),  market;  MCZ  1037  (1);  MCZ  25741  (1);  MNHN 
6369  (holotype  of  Malthaea  longirostris  Cuvier  and  Valen- 
ciennes): ZMUC  18  (1).  Espirito  Santo:  CAS-SU  59612  (1) 
and  CAS-SU  59614  (1),  Enseada,  Vitoria;  CAS-SU  59615  (1), 
Vitoria.  Rio  de  Janeiro:  MCZ  11755  (1)  and  MCZ  11756  (2), 
Rio  de  Janeiro.  Sao  Paulo:  CAS-SU  59610  (1)  and  CAS-SU 
59613  (1),  Ponta  da  Praia,  Santos;  CAS-SU  59611  (1),  Ponta 
do  Rabo  Azedo,  1 13  km  NE  Sao  Paulo. 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


251 


FIGURE  12.  Variation  in  Ogcocephalus  nasutus  (Cuvier  and  Valenciennes).  (A)  Specimen  from  off  Surinam,  FMNH  87034, 
135.0  mm  SL,  and  (B)  specimen  (juvenile)  from  Caribbean  off  Venezuela,  MCZ  41510,  84.0  mm  SL,  both  showing  common 
pattern  of  clusters  of  spots  on  shoulders.  (C)  Specimen  from  northern  Brazil,  FMNH  66386,  130.0  mm  SL,  showing  uncommon 
condition  of  clusters  of  spots  extending  from  eyes  backwards  to  axillae,  reminiscent  of  O.  vespertilio.  (D)  From  Honduras, 
CAS  28520,  148.5  mm  SL.  (E)  and  (F)  From  Surinam,  CAS-SU  51208,  130.5  mm  SL,  and  FMNH  64900,  203.0  mm  SL.  (G) 
From  Nicaragua,  CAS  28494,  205.0  mm  SL.  Extremely  short  rostrum  (D),  bent  rostrum  (F),  and  extremely  long  rostrum  (G) 
are  unusual,  but  all  occur  in  both  Central  American  and  South  American  material. 


Ogcocephalus  nasutus  (Cuvier  and  Valenciennes) 

(Figures  12  and  13) 

[?]  Rana  piscatrix  SEBA,  1734:pl.  74,  fig.  2  [Curacao;  see  com- 
ment after  synonymy]. 

Lophius  vespertilio  (non  Linnaeus):  BLOCH  1787:8,  pi.  110 
[description];  PARRA  1787: pi.  4  [not  seen;  a  hand-drawn 
copy  by  Senorita  Poey  was  consulted:  Cuba];  CUVIER 
1816:311  [based  on  Bloch  1787]. 

[?]  Lophius  rostratus  SHAW,  1804:383  [description;  nomen 
dubium;  see  comment  following  synonymy]. 

[?]  Malthe  nasuta  CUVIER,  1829:252  [nomen  dubium,  based 
on  Seba  1734;  see  comment  following  synonymy]. 

Malthea  vespertilio  (non  Linnaeus):  CUVIER  AND  VALEN- 
CIENNES 1837:440  ["Porto-Rico,  Martinique,  Terre-Neuve"; 
description]. 


Malthaea  nasuta  CUVIER  AND  VALENCIENNES,  1837:452 
[Martinique,  "Saint-Domingue,  New- York"]. 

Ogcocephalus  vespertilio  (non  Linnaeus):  JORDAN  AND  Ev- 
ERMANN  1898:2737  ["Havanna,  Cuba";  description];  MEEK 
AND  HILDEBRAND  1928:1017  [Porto  Bello,  Panama; 
description];  BEEBE  AND  TEE-VAN  1928:272  [Port-au-Prince 
Bay,  Haiti;  key;  field  characters;  color  in  life];  HERRE 
1942:305  [Bridgetown,  Barbados];  MARTIN  S.  1956: 142  [Los 
Roques  off  coast  of  Venezuela;  description];  [?]  LOWE 
(McCoNNELL)  1962:680  [British  Guiana  (Guyana);  partial 
description;  ecology];  [?]  CERVIGON  M.  1966:870  [Vene- 
zuela; description;  key];  [?]  DAHL  1971:317  [Colombia; 
key]. 

Ogcocephalus  nasutus:  BEEBE  AND  TEE-VAN  1928:273  [Port- 
au-Prince  Bay,  Haiti;  key;  field  characters;  color  in  life]; 
BRADBURY  1967:417  [listed];  RANDALL  1967:825  [West  In- 
dies; food  habits];  1968:295  [photograph;  description]; 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


FIGURE  13.  Variation  in  Ogcocephalus  nasutus  (continued  from  Fig.  12).  (A)  From  Belize,  CAS  42579,  182.0  mm  SL,  and 
(B)  from  the  Bahamas,  ANSP  90674,  187.0  mm  SL,  both  lacking  color  pattern  except  for  few  dim  ocellated  spots  on  disk  of 
Belize  specimen.  In  spite  of  absence  of  color  pattern  on  body,  striped  and  spotted  patterns  on  face  common  in  specimens  from 
West  Indies  and  Central  America:  (D)  from  Panama,  MCZ  11768,  112.0  mm  SL;  (E)  from  Belize,  same  specimen  as  (A);  (F) 
from  the  Bahamas,  UF  24185,  108.0  mm  SL.  (D)  Also  shows  exaggeratedly  spiny  bucklers  occasionally  seen  among  specimens 
from  West  Indies  and  Central  America;  (F),  a  less  pronounced  example  of  spiny  squamation.  (C)  From  the  Bahamas,  ANSP 
90670,  170.0  mm  SL,  showing  clusters  of  spots  frequently  seen  in  West  Indies  material;  however,  this  specimen  also  has  square- 
browed  conformation  of  head  characteristic  of  O.  cubifrons  and  is  intermediate  between  cubifrons  and  nasutus  in  other 
characters.  (G)  From  the  Bahamas,  ANSP  90671,  162.0  mm  SL,  also  intermediate  between  cubifrons  and  nasutus,  with 
extremely  dark  coloration  sometimes  seen  in  West  Indies  material,  but  photographed  while  wet  and  intensely  highlighted; 
compare  conformation  of  rostrum  with  Figure  23G. 


BOHLKE  AND  CHAPLIN  1968:728  [description;  bathymetric 

range;  habits;  key;  figure]. 
Oncocephalus  vespertilio  (non  Linnaeus):  FOWLER  1947: 14, 

fig.  9  [Nassau  or  Hog  I.,  Bahama  Is.;  description]. 
Ogcocephalus  radiatus:  BREDER  1949:95,  pi.  10,  figs.  \-4  [Bi- 

mini,  Bahama  Is.;  behavior;  photographs];  DAHL  1971:317 

[Colombia;  key;  figs.]. 

The  name  Lophius  rostratus  Shaw,  1804,  is 
based  upon  a  textbook  compilation  drawn  from 
Linnaeus  together  with  the  pre-Linnaean  au- 
thors Seba  (1734),  Browne  (1756),  and  Marcgrave 
(1648).  The  description  in  Marcgrave  of  a  fish 
from  Brazil  gives  proportions  of  the  rostrum  in- 
dicating it  was  the  same  species  Linnaeus  later 
described  and  named  vespertilio.  Linnaeus  used 


the  term  "rostratus"  (beaked)  in  describing  ves- 
pertilio, suggesting  that  Shaw's  name  derives 
from  Linnaeus's  description.  In  any  case,  ros- 
tratus Shaw  in  part  refers  to  the  long-nosed  Bra- 
zilian form  described  by  Marcgrave  and  Lin- 
naeus. However,  the  other  two  authors  to  whom 
Shaw  referred,  Seba  and  Browne,  described  and 
figured  specimens  said  to  be  from  Curacao  and 
Jamaica,  respectively;  the  localities  and  the  fig- 
ures suggest  nasutus  but  do  not  limit  the  pos- 
sibilities to  nasutus.  The  illustration  in  Shaw 
also  appears  to  depict  nasutus  because  it  shows 
a  rostrum  of  only  moderate  proportions,  but 
apart  from  that  the  illustration  is  highly  stylized 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


253 


and  difficult  to  interpret.  Finally,  the  range  given 
by  Shaw,  "South- American  seas,"  is  too  gen- 
eral to  apply  to  any  particular  species.  As  I  am 
unable  to  determine  Shaw's  Lophius  rostratus 
either  from  his  sources  or  from  his  description, 
I  hereby  reject  the  name  Lophius  rostratus 
Shaw  as  a  nomen  dubium. 

The  name  Malthe  nasuta  Cuvier,  1829,  was 
published  in  a  footnote  with  no  other  designation 
than  a  figure  in  Seba  (1734)  labeled  Rana  pis- 
catrix.  This  stylized  drawing  may  represent  the 
Antillean-Caribbean  form  of  the  species  of 
Ogcocephalus  under  consideration  here,  but  it 
is  a  very  generalized  figure  and  could  be  a  rep- 
resentation of  any  of  several  other  species.  Be- 
cause it  is  impossible  to  establish  the  identity  of 
the  figure  upon  which  the  name  Malthe  nasuta 
Cuvier,  1829,  was  based,  the  original  description 
of  Ogcocephalus  nasutus  is  here  considered  to 
be  that  of  Cuvier  and  Valenciennes,  1837. 

COMPARISONS  AND  DIAGNOSIS. — At  the  be- 
ginning of  this  study,  only  specimens  from  the 
West  Indies  and  the  Venezuela-Guiana  coast  of 
South  America  were  available  for  examination. 
These  specimens  showed  some  differences  in 
color  pattern  and  sometimes  in  squamation. 
Specimens  from  the  West  Indies  included  many 
with  relatively  dark  overall  color  and  reticulated 
markings  around  the  gill  pores,  in  the  axillae, 
and  on  the  lateral  sides  of  the  tail;  these  some- 
times had  jagged  clusters  of  spinules  at  the  tips 
of  bucklers  that  made  the  body  very  rough  (Fig. 
13D  shows  this  condition).  Specimens  from  the 
northern  coast  of  South  America  were  a  lighter, 
more  uniform  brown  in  color  with  less  jagged 
bucklers;  these  generally  were  from  depths  of  30 
m  or  more,  but  no  comparisons  with  the  West 
Indies  specimens  could  be  made  on  this  point, 
as  the  latter  usually  lacked  bathymetric  data. 

Eventually,  specimens  from  the  Caribbean 
coast  of  Central  America  became  available. 
They  generally  came  from  offshore  in  depths 
over  30  m,  but  some  were  from  shallower  water. 
These  Central  American  specimens  resembled 
the  South  American  specimens  in  color  and 
squamation,  their  integument  relatively  pale  and 
smooth,  but  the  dark,  rough-skinned  variants 
occur  here  too,  known  from  older  museum  spec- 
imens that  also  lack  bathymetric  data.  In  any 
case,  the  three  groups  of  specimens  were  treated 
separately,  so  tables  of  data  in  this  report  show 
three  sets  of  data  for  O.  nasutus,  which  are  des- 


ignated as  West  Indies  nasutus,  Central  Amer- 
ican nasutus,  and  South  American  nasutus. 

Besides  varying  in  color  pattern,  O.  nasutus 
is  more  variable  than  other  species  in  the  genus 
in  lateral-line  scale  counts  and  in  rostrum  pro- 
portions, features  relied  upon  in  this  study  to 
characterize  species.  Apparently,  nasutus  also 
has  a  greater  bathymetric  range  than  any  of  its 
congeners,  occurring  inshore  as  well  as  offshore 
to  depths  of  approximately  200  m  in  parts  of  its 
range.  Perhaps  bathymetric  variations  will  even- 
tually help  explain  morphological  variation  in 
this  form. 

One  of  the  largest  species  of  Ogcocephalus 
(Table  1),  nasutus  most  resembles  cubifrons  to 
the  north  of  its  range  and  vespertilio  to  the 
south.  At  this  time  it  appears  that  nasutus  is 
allopatric  with  the  Brazilian  species  vespertilio, 
but  this  assertion  needs  verification.  In  the  north 
nasutus  occurs  with  cubifrons  off  southeastern 
Florida  and  in  the  western  Bahamas;  whether  or 
not  it  is  entirely  distinct  from  cubifrons  in  the 
western  Bahamas  is  in  doubt  because  a  few 
specimens  which  are  intermediate  between  na- 
sutus and  cubifrons  have  been  taken  there.  O. 
nasutus  differs  from  cubifrons  in  having  a  longer 
jaw  (Fig.  14)  and  a  narrower  interorbital  distance 
(Fig.  15),  a  rostrum  longer  on  the  average  but 
more  variable  (Fig.  10),  and  higher  modal  num- 
ber of  lateral-line  scales  in  the  cheek  series  (9 
instead  of  8  as  in  cubifrons,  Table  5).  Color  pat- 
tern also  assists  in  separating  nasutus  from  cub- 
ifrons: clusters  of  dark  spots  occur  on  the  face, 
disk,  and  tail  in  both  species,  but  nasutus  lacks 
the  bright  pattern  of  spots  on  the  pectoral  fins 
seen  in  cubifrons.  From  vespertilio,  nasutus  dif- 
fers in  usually  having  a  shorter  rostrum  (Fig. 
10),  a  narrower  interorbital  distance  (Fig.  11), 
and  a  lower  modal  count  of  pectoral  rays  (12-13 
in  nasutus,  14  in  vespertilio,  Table  3). 

Ogcocephalus  nasutus  also  resembles  panto- 
stictus  and  rostellum  in  general  appearance. 
Compared  to  pantostictus,  which  shows  allo- 
metric  growth  of  the  rostrum,  the  rostrum  in 
nasutus  is  variable  but  generally  longer  than  in 
pantostictus  (Fig.  10).  O.  nasutus  also  differs 
from  pantostictus  in  having  a  lower  modal  num- 
ber of  lateral-line  scales  in  the  subopercular  se- 
ries (8  compared  to  9,  Table  6),  and  nasutus  has 
a  pattern  of  spot  clusters  or  may  lack  spots  en- 
tirely, while  pantostictus  has  a  pattern  resem- 
bling polkadots  covering  the  entire  dorsal  sur- 


254 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


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FIGURE  14.     Comparisons  between  Ogcocephalus  nasutus  (only  specimens  from  West  Indies),  O.  cubifrons,  and  O.  pan- 
tostictus  to  show  that  jaw  is  short  in  O.  cubifrons  relative  to  width  of  interorbital. 


face  of  the  body.  O.  nasutus  differs  from 
rostellum  in  having  a  longer  rostrum  (Fig.  10), 
a  higher  modal  number  of  lateral-line  scales  in 
the  cheek  series  (9  compared  to  8  as  found  in 
rostellum,  Table  5),  and  a  relatively  longer  jaw 
(Fig.  16) 

Ogcocephalus  parvus  and  O.  pumilus  occur 
within  the  range  of  nasutus,  but  both  are  smaller 
species,  reaching  only  85  mm  in  standard  length. 
Both  differ  further  from  nasutus  in  some  meris- 
tic  counts:  pectoral  rays  usually  12-13  in  nasu- 
tus, 10-12  in  parvus  and  pumilus;  lateral-line 
scales  in  the  cheek  series  9  in  nasutus,  8  in  the 
other  two  (Tables  3  and  5). 

The  range  of  O.  nasutus  slightly  overlaps  that 
of  corniger  in  the  vicinity  of  southeastern  Flor- 
ida and  the  western  Bahamas;  however,  corni- 
ger is  distinguished  by  its  color  pattern  of  close- 
set  pale  spots  on  a  darker  background  evenly 
covering  the  entire  dorsal  surface,  while  nasutus 
has  spot  clusters  on  a  lighter  background  or  no 


spots  at  all.  O.  nasutus  and  O.  corniger  also 
differ  in  some  meristic  counts:  pectoral  rays 
usually  12-13  in  nasutus,  10-12  in  corniger;  lat- 
eral-line scales  in  the  cheek  series  9  in  nasutus, 
8  in  corniger  (Tables  3  and  5). 

The  southern  portion  of  the  range  of  nasutus 
overlaps  the  range  of  notatus,  but  the  two  are 
distinguished  by  the  relative  length  of  the  disk 
margin  (longer  in  nasutus,  Fig.  7)  and  width  of 
the  mouth  (narrower  in  nasutus,  Fig.  6)  as  well 
as  the  lateral-line  scale  count  of  3  in  the  pre- 
maxillary  series  in  nasutus  (4  in  notatus)  and  9 
in  the  cheek  series  (10  in  notatus). 

DESCRIPTION. — Counts  and  measurements 
based  on  106  specimens  as  follows:  40  speci- 
mens, 26.2-232.0  mm  SL  from  the  West  Indies; 
25  specimens  42.7-223.0  mm  SL  from  the  coast 
of  Central  America  and  Yucatan;  41  specimens, 
29.0-233.5  mm  SL  from  the  coast  of  South 
America. 

Counts.   Counts  given  in  Tables  2-6.  A 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


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FIGURE  15.     Comparison  between  Ogcocephalus  cubifrons  and  O.  nasutus  to  show  that  interorbital  is  widest  in  O.  cubifrons 
relative  to  length  of  disk  margin. 


marked  difference  in  pectoral  count  exists  be- 
tween Antilles- West  Indian  and  Central  Amer- 
ican specimens  on  the  one  hand  and  South 
American  specimens  on  the  other,  most  fre- 
quently 12  in  Antilles- West  Indian  and  Central 
American  specimens,  usually  13  in  South  Amer- 
ican specimens.  A  small  cline  is  formed  between 
these  two  groups  of  nasutus  and  vespertilio, 
which  most  frequently  has  14  pectoral  rays,  but 
clinal  variation  is  not  found  in  other  counts.  O. 
nasutus  shows  a  wider  range  of  variation  with 
a  higher  degree  of  asymmetry  in  counts  of  lat- 
eral-line scales  in  the  subopercular  series  than 
other  species  in  the  genus,  but  resembles  most 
other  species  in  having  a  relatively  narrow  range 
of  variation  in  the  count  for  the  cheek  series  of 
lateral-line  scales  (modally  9,  a  feature  in  which 
nasutus  resembles  vespertilio).  O.  nasutus  has 
a  relatively  high  count  for  lateral-line  scales, 
perhaps  associated  with  the  large  size  of  the 
species,  since  other  large  species  (vespertilio, 
pantostictus,  and  cubifrons)  also  have  high  lat- 
eral-line scale  counts. 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses. 

Length  of  disk  margin  intermediate  for  the 
range  of  variation  in  the  genus,  2.2(2.0-2.4)  in 
SL.  Tail  moderately  wide  to  narrow,  its  width 
2.0(1.7-2.7)  in  length  of  disk  margin;  depth  of 
caudal  peduncle  quite  variable  but  averaging  rel- 
atively deep,  2.8(2.2-3.5)  in  head  depth.  Ros- 


trum extremely  variable  in  length  and  contour. 
It  may  be  as  short  as  the  rostrum  of  examples 
of  cubifrons  of  comparable  size  (especially  true 
among  Bahaman  and  western  Caribbean  speci- 
mens) or  as  long  as  that  of  vespertilio.  It  may 
be  thick-based  and  conical  or  slender  and  finger- 
shaped.  Usually  it  is  straight,  either  horizontal 
or  tilted  upward,  but  a  few  specimens  have  it 
crooked  with  its  tip  bent  sharply  upward  or 
drooping  downward  (Fig.  12,  13).  Length  of  ros- 
trum 3.7(1.9-5.7)  in  length  of  disk  margin.  Ap- 
erture of  illicial  cavity  subtriangular,  higher  than 
wide.  Cranium  rising  steeply  above  disk,  head 
depth  2.0(1.7-2.2)  in  length  of  disk  margin. 
Width  of  cranium  3.4(2.5-4.0)  in  length  of  disk 
margin.  The  following  in  width  of  cranium:  eye 
1.5(1.3-1.9),  lateral  ethmoid  width  1.7(1.4-2.0). 
Interorbital  variable,  from  slightly  concave  to 
flat  or  even  a  little  convex  anteriorly  where  the 
interorbital  contours  grade  into  the  roundness  of 
the  rostrum.  Width  of  interorbital  2.1(1.7-2.7)  in 
width  of  cranium,  3.6(2.8-4.4)  in  head  depth. 
Mouth  moderate,  its  width  1.6(1.4-1.8)  in  head 
depth,  3.2(2.7-3.8)  in  length  of  disk  margin;  jaw 
2.3(2.1-2.5)  in  head  depth.  Upper  lip  fleshy,  of 
an  even  width  for  its  entire  length;  lower  lip 
fleshy,  swollen  medially.  Dorsal  fin  4.4(2.9-5.6) 
in  disk  margin;  anal  fin  3.0(2.4-3.9)  in  disk  mar- 
gin. 

Integument.  Dermal  cirri  sometimes  present. 
Large  bucklers  prominent,  arranged  as  in  ge- 
neric description.  All  fins  with  tiny  tubercles. 


256 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


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FIGURE  16.     Comparison  between  Ogcocephalus  nasutus  (only  specimens  from  West  Indies)  and  O.  rostellum  to  show  that 
jaw  is  long  in  O.  nasutus  relative  to  width  of  cranium. 


On  pectorals,  ventrals,  and  anal,  tubercles  are 
variable  in  distribution,  sometimes  restricted  to 
bases  of  fins,  sometimes  running  out  along  rays 
for  half  their  length.  Dorsal  fin  with  tubercles  on 
anterior  edge  of  first  ray,  sometimes  running  out 
along  ray  nearly  to  its  tip.  Caudal  as  in  generic 
description.  Skin  covering  eyeballs  bearing  tiny 
tubercles  with  rim  around  the  cornea  beaded  by 
a  row  of  small  bucklers.  Pectoral  membranes 
thick,  opaque,  ventral  surface  of  rays  with 
fleshy  pads. 

Color  in  preservative.  Specimens  from  West 
Indies  often  very  darkly  pigmented,  but  speci- 
mens from  off  coast  of  South  America  and  Cen- 
tral America  usually  have  dorsal  surface  of  body 
tan  or  medium  brown,  with  ventral  surface  of 
body  usually  paler  than  dorsal  surface.  As  point- 
ed out  elsewhere,  it  is  likely  that  this  color  vari- 
ation is  correlated  with  depth  and  not  geograph- 
ical locality;  a  corroborative  observation  is  that 
several  specimens  from  Central  America  from 
presumed  shallow  water  (exact  depths  not  giv- 
en) were  relatively  darkly  pigmented. 

Roundish  dark  spots  often  present  clustered 


in  a  patch  on  either  shoulder.  Markings  on  face, 
lateral  walls  of  tail,  and  in  axillae  tend  to  be 
reticula  of  pale  lines  between  dark  irregular 
patches.  Specimens  from  West  Indies  some- 
times have  dark  stripes  with  narrow  light  bor- 
ders on  face  and  iris  which  form  a  radiating  pat- 
tern around  pupil.  Ventral  surface  pale  to  dark 
gray  in  West  Indian  specimens,  the  chin  often 
particularly  dark;  ventral  surface  uniformly  pale 
in  South  American  and  Central  American  spec- 
imens. Lips  in  West  Indian  specimens  usually 
darkly  pigmented  in  dark  specimens,  but  oth- 
erwise lips  pale.  Rostrum  generally  with  a  dif- 
fuse black  subterminal  ring.  Iris  black  or  marked 
with  an  extension  of  color  markings  efface.  Pec- 
torals of  West  Indian  specimens  generally  dark 
to  nearly  black,  always  with  a  black  margin  even 
when  fin  pale  basally;  pectorals  of  South  Amer- 
ican and  Central  American  specimens  usually 
pale  dusky  to  medium  gray  or  tan,  shading  to  a 
darker  margin.  Dorsal  variable,  pale  to  dark, 
ordinarily  without  markings.  Ventrals  and  anal 
the  same  ground  color  as  ventral  surface  of 
body;  in  dark  specimens  the  ventrals  and  anal 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


257 


®  nasutus 
•  cubifrons 
©  pantostictus 
o  rostellum 


FIGURE  17.     Localities  for  study  material  of  Ogcocephalus  nasutus,  O.  cubifrons,  O.  pantostictus,  and  O.  rostellum.  Scale 
of  map  too  small  to  show  that  O.  cubifrons  and  O.  pantostictus  are  parapatric  in  northern  Gulf  of  Mexico;  see  text  for  localities. 


may  have  dark  tips,  and  small  specimens  nearly 
always  have  anal  black-tipped.  Caudal  fin  as  in 
generic  description. 

Color  in  life.  Kodachrome  transparencies 
made  of  a  freshly  captured  specimen  from  off 
coast  of  South  America  show  dorsal  surface  of 
body  uniformly  grayish  olive-green,  a  mudlike 
color,  with  conspicuous  clusters  of  black  spots 
on  either  shoulder.  Pectorals  and  caudal  the 
same  greenish  shade  basally,  shading  to  black 
margins.  Lips,  ventral  surface  of  body,  and  anal 
fins  rosy  tan. 

Breder  (1949)  writes  of  a  specimen  captured 
from  off  Bimini  and  kept  in  an  aquarium,  "For 
the  most  part  the  fish  remained  an  intense  black, 
so  much  so  that  it  was  only  with  difficulty  that 
any  of  the  ordinarily  visible  fish  structures  could 
be  seen.  Thus  the  pupil,  iris,  and  skin  about  the 
eye  were  so  much  the  same  that  none  of  the 
details  could  ordinarily  be  seen  at  all.  The  inside 
of  the  mouth  and  the  membranes  shown  when 
'yawning'  were  at  all  times  a  rich  mahogany  and 
very  contrasting  to  the  entirely  black  fish.  At 
night  on  a  few  occasions  the  iris  became  ma- 
hogany colored,  and  vague  lighter  bands  ap- 
peared across  the  back." 

Perhaps  color  variation  is  governed  to  some 
extent  by  variations  in  the  amount  of  ambient 
light,  with  dark  specimens  coming  from  shallow, 


well-lighted  waters  and  lighter  colored  speci- 
mens coming  from  deeper  water.  Dr.  John  E. 
Randall  (personal  communication)  says  of  a 
specimen  164  mm  SL  which  was  taken  in  water 
about  21  m  deep  in  Reef  Bay,  St.  John,  Virgin 
Islands,  "The  fish  was  brown  on  top  with  no 
spots.  There  were  some  short  white  lines  near 
the  gill  opening  and  around  the  ventral  half  of 
the  eye  and  snout  below  the  rostral  prominence. 
The  mouth  was  bright  red.  The  ventral  part  of 
the  fish  was  red." 

Dr.  Bruce  B.  Collette  (personal  communica- 
tion) noted  that  a  specimen  he  took  from  water 
1-1.5  m  deep  off  the  Isle  of  Pines  had  lips  that 
were  brilliant  vermillion. 

DISTRIBUTION  (Fig.  17). — Material  used  for 
counts  and  measurements  in  this  study  came 
from  the  West  Indies  and  the  coasts  of  Central 
and  northern  South  America  to  the  mouth  of  the 
Amazon  River.  However,  during  the  final  prep- 
aration of  this  report,  the  following  specimens 
were  seen  which  extend  the  range  to  southeast- 
ern Florida:  UMML  16813,  1  specimen,  bridge 
at  Bear  Cut,  Dade  Co.,  and  UMML  33252,  1 
specimen,  Bear  Cut  at  Virginia  Key,  Dade  Co. 
Bathy metric  range:  shore  to  275  m. 

REMARKS. — Early  in  this  study  I  examined  the 
type  material  in  the  Museum  National  d'Histoire 
Naturelle,  Paris,  which  consisted  of  the  follow- 


258 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


ing  three  specimens:  A. 4665,  Martinique,  Ke- 
raudren;  A. 4765,  Saint  Domingue,  Ricord  (Pi- 
chon);  and  A. 4766,  New  York,  Milbert.  The 
Martinique  specimen,  a  dried  example  mounted 
on  a  wooden  stand  and  about  200  mm  SL,  was 
difficult  to  perform  counts  and  measurements 
on,  but  had  a  relatively  short  rostrum,  wide 
mouth,  and  narrow  interorbital.  The  Saint  Do- 
mingue specimen,  192.5  mm  SL,  had  an  ex- 
tremely short  rostrum,  wide  mouth,  and  wide 
interorbital.  The  New  York  specimen,  34.0  mm 
SL,  had  a  relatively  long,  slender  rostrum,  but 
the  specimen  is  too  small  for  other  proportions 
to  be  useful.  This  specimen  is  also  problematical 
because  the  locality  is  outside  the  known  range 
of  nasutus  as  understood  here.  Although  a  stray 
occurrence  off  New  York  is  certainly  possible, 
it  is  also  possible  that  the  original  locality  infor- 
mation is  wrong.  None  of  the  specimens  had 
discernible  markings. 

Recognizing  now  that  nasutus  is  variable  and 
difficult  to  characterize,  and  that  setting  its  lim- 
its is  yet  to  be  accomplished,  I  cannot  assert  that 
the  three  types  are  conspecific.  To  compound 
the  difficulty,  I  recently  checked  the  status  of 
these  specimens  and  was  informed  by  Dr.  M.- 
L.  Bauchot  (personal  communication)  that  spec- 
imen A. 4665  (the  Martinique  specimen)  unfor- 
tunately has  disappeared  from  the  collection. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  For  data  from  OREGON  stations  (  =  O),  refer 
to  Bullis  and  Thompson  (1965). 

West  Indies:  Bahamas:  AMNH  18025  (1);  ANSP  72336  (1), 
Nassau  and  Hog  I.;  ANSP  90669  (1)  and  ANSP  90673  (1),  N 
shore  of  Hog  I.;  ANSP  90672  (1)  and  90674  (1),  S  shore  of 
Hog  I.;  MCZ  11767  (2),  Nassau;  UF  24185  (1),  SILVER  BAY 
sta.  2464,  23°34'N,  79°05'W,  274  m.  Cuba:  ANSP  89011  (1), 
Havana;  CAS-SU  3096  (1),  Havana;  CAS  27215  (1)  and  CAS 
42578  (1),  Havana;  MCZ  11758-11760  (5);  MCZ  11763  (3); 
MCZ  350%  (1),  Guantanamo  Bay:  USNM  13051  (1).  Jamaica: 
CAS-SU  4744  (3).  Haiti:  USNM  132575  (1),  Port-au-Prince; 
USNM  178027  (1).  Dominican  Republic:  AMNH  8297  (2)  and 
AMNH  17462(1);  ZMUC  10(1)  and  ZMUC  P9229(l),  Puerto 
Plata.  Puerto  Rico:  FMNH  64842  (1),  O-2626;  CAS  23918  (1), 
O-2668.  Virgin  Islands:  FMNH  53947  (1),  St.  Croix;  UMML 
6821  (1),  Wend  Reef  Bay,  St.  John;  USNM  116432(1),  Shoy 
Lagoon,  St.  Croix;  ZMUC  5  (1),  St.  Croix.  Lesser  Antilles: 
ANSP  10622-10623  (2),  St.  Martin;  MCZ  27332  (1),  "St. 
Bart."  [  =  Saint  Barthelemy?];  CAS-SU  39893  (1),  Bridge- 
town, Barbados;  FMNH  64847  (1),  O-2631. 

Central  America:  British  Honduras:  AMNH  8209  (1),  Porta 
Cortez:  CAS  28495  (2),  O-6405,  17°18'N,  88°01'W,  183  m: 
CAS  42579  (2),  UNDAUNTED  cr.  6703,  sta.  72,  17°12'N, 
88°11.2'W,  18-20  m;  USNM  37123  (1),  off  Cozumel;  BMNH 
1927.9.16.18-22  (4),  Watercey  I.  Honduras:  CAS  28520  (1), 
HURRICANE,  15°48'N,  83°54'W,  24  m:  CAS  28521-22  (4), 
SHADY  LADY,  15°49.5'N,  83°44'W,  31  m;  CAS  28523  (2), 


SHADY  LADY,  15°54'N,  83°40'W,  37  m;  FMNH  64329  (2), 
O-1934;  FMNH  64330  (4),  O-1937.  Nicaragua:  CAS  28494  (2), 
O-6426,  12°56'N,  82021'W,  190  m.  Panama:  MCZ  11768  (1), 
Isthmus  of  Darien;  USNM  81728  (1),  Porto  Bello. 

South  America:  Venezuela:  CAS  28527  (1),  CALAMAR  sta. 
502,  H°15'N,  65°15'W,  82-104  m;  FMNH  64900  (4),  O-2348; 
MCZ  41510  (4),  ATLANTIS  sta.  5599,  10°19'N,  65°32.5'W,  64 
m.  Guyana:  CAS  28525  (1),  CALAMAR  sta.  425,  95-101  m; 
FMNH  64894  (1),  O-2251;  FMNH  65950  (2),  O-2249;  FMNH 
65960  (4),  O-2231;  FMNH  65961  (2),  O-2233;  FMNH  65962 
(2),  O-2254;  FMNH  66386  (1),  O-1996;  UF  24184  (1),  O-2241. 
Surinam:  CAS  23915  (1),  O-2279;  CAS  23917  (1),  COQUETTE 
sta.  178,  6°19'N,  55°50.5'W,  27  m;  CAS-SU  51208  (1),  CO- 
QUETTE sta.  30,  6°49.5'N,  54°54'W,  48  m;  CAS-SU  51211  (1), 
COQUETTE  sta.  213,  6°46'N,  54°29.5'W,  44  m;  FMNH  648% 
(1),  O-2275;  FMNH  64898  (8),  O-2329;  FMNH  65959  (1),  O- 
2016;  FMNH  65%3  (1),  O-2266;  MCZ  45076  (1),  COQUETTE 
sta.  182,  6°15'N,  55°54'W,  24  m;  UMML  4012  (1),  O-2268. 
French  Guiana:  CAS  23916  (1),  O-2043;  FMNH  64889  (1),  O- 
2035;  FMNH  64890  (1),  O-2036;  FMNH  64897  (1),  O-2303; 
FMNH  65964  (1),  O-2301.  Brazil:  MCZ  45075  (4),  O-2074. 

Undetermined  material.  Seven  specimens  from  the  Bahama 
Islands  are  intermediate  in  color  pattern  between  nasutus  and 
cubifrons,  and  also  cannot  be  diagnosed  with  certainty  on  the 
basis  of  morphometric  characters;  these  are  ANSP  72335  (1), 
Hog  I.,  ANSP  90670-90671  (2),  south  shore  of  Hog  I.,  and 
FMNH  64139  (4),  Grand  Bahama  I. 

Ogcocephalus  cubifrons  (Richardson) 

(Figures  18  and  22) 

[?]  Lophius  radiatus  MITCHILL,  1818:326  ["Strait  of  Baha- 
ma": nomen  dubium:  see  comment  after  synonymy]. 

Lophius  (Malthe)  cubifrons  RICHARDSON,  1836:103,  pi.  96 
[Labrador  (see  comment  after  synonymy);  holotype  BMNH 
1855.9.19.848]. 

Malthaea  cubifrons:  CUVIER  AND  VALENCIENNES  1837:452 
[based  on  Richardson  1836;  cubifrons  placed  in  synonymy 
of  mis u tits]. 

Malthea  nasuta  (non  Cuvier  and  Valenciennes):  STORER 
1846:131  [synonymy:  partial  description:  range,  compiled]. 

Malthe  cubifrons:  GUNTHER  1861:203  [partial  description  of 
holotype];  GILL  1873:14  [Labrador;  name  only]:  GOODE 
AND  BEAN  1879:121  [west  Florida:  fin  formulae]:  1882:235 
[Gulf  of  Mexico:  name  only];  JORDAN  AND  GILBERT 
1882:850  [coasts  of  Florida;  partial  description]. 

Malthea  radiala:  GILL  1862:47  [name  only:  based  on  Malthea 
nasuta  of  Storer  1846]. 

Malthe  vexpertilio  (non  Linnaeus):  GILL  1873:14  [Labrador, 
compiled];  JORDAN  AND  SWAIN  1885:234  [Cedar  Keys, 
Florida:  partial  description;  cubifrons  and  nasutus  referred 
to  synonymy  of  vespertilio]:  HENSHALL  1895:220  [Tampa, 
Florida];  JORDAN  1899:172  [name  listed]. 

Malthe  respertilio  nasuta  (non  Cuvier  and  Valenciennes): 
JORDAN  AND  GILBERT  1882:850  [range:  South  Atlantic  coast 
of  United  States:  partial  description]. 

Malthe  vespertilio  radiata:  JORDAN  1885a:927  (part)  [variety 
cubifrons  referred  to  the  synonymy  of  radiatus]:  1885b:  144 
[Key  West,  Florida;  partial  description):  1885c:45  [Egmont 
Key,  Florida]. 

Malthe  radiata:  HENSHALL  1891:383  [southern  Florida];  1895: 
220  [Tampa,  Florida;  partial  description]. 

Onchocephalus  radiatus:  GOODE  AND  BEAN  1896:498  [Pen- 
sacola,  Florida:  description:  notes  on  holotype  of  c ubifrons 
Richardson,  1836]. 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


259 


FIGURE  18.  Ogcocephalus  cubifrons  (Richardson)  from  west  coast  of  Florida  showing  interorbital  width  increases  with  size 
and  rostrum  length  decreases.  Left  to  right:  78.0  mm  SL,  FSBC  342,  64-81  km  NE  Dry  Tortugas;  98.0  mm  SL,  UF  2730,  off 
Pensacola;  194.0  mm  SL,  FSBC  1369,  off  Don-ce-Sar,  Pinellas  County. 


Ogcocephalus  vespertllio  (non  Linnaeus):  JORDAN  AND  Ev- 

ERMANN  1896:511  (part)  [list,  compiled]:  EVERMANN  AND 
KENDALL  1900:99  (part)  [locality  records,  compiled]:  GUN- 
TER,  WILLIAMS,  DAVIS,  AND  SMITH  1948:314  [Fort  Myers, 
Florida:  mortality]. 

Ogcocephalus  radiatus:  JORDAN  AND  EVERMANN  1898:2738 
[Cedar  Key,  Florida;  description:  range;  radiatus  (=cubi- 
frons)  removed  from  synonymy  of  vespertilio]:  EVERMANN 
AND  KENDALL  1900:99  (part)  [Key  West,  Florida;  other  lo- 
calities compiled];  FOWLER  1906:108  [Florida  Keys:  color 
in  life:  behavior:  figure]:  1915:251  [Palm  Beach,  Florida]: 
BREDER  1929:303  [Florida];  JORDAN,  EVERMANN,  AND 
CLARK  1930:507  [compiled]:  STOREY  AND  GUDGER  1936:648 
[mortality]:  GUNTER,  WILLIAMS,  DAVIS,  AND  SMITH 
1948:312  [Englewood,  Florida:  mortality]:  BRIGGS  1958:301 
[range,  compiled):  BOHLKE  AND  CHAPLIN  1968:727  [Baha- 
ma Is.;  description:  key:  figure];  WALLS  1975:117  [partial 
description;  color  photograph). 

Oncoceplialiis  nidi  at  us:  FOWLER  1926:254  [Captiva  Pass, 
Florida]:  1945:335  (part)  [Sanibel  I.,  Florida]. 

Ogcocephalus  cubifrons:  LONGLEY  AND  HILDEBRAND 
1941:311-312,  pi.  33,  fig.  2  [Tortugas,  Florida:  key: 
description);  HILDEBRAND  1955:219  ([Campeche  Banks, 
Yucatan:  partial  description):  SPRINGER  AND  MCERLEAN 
1962:41  [south  Florida:  listed):  MOE  AND  MARTIN  1965:147 
[off  Pinellas  Co.,  Florida:  ecological  survey:  listed):  MOE 
et  al.  1966:120  [Florida:  list  of  museum  specimens):  BRAD- 
BURY 1967:417  [listed]:  STARK  1968:33  [Alligator  Reef,  Flor- 
ida: composition  of  reef  fauna). 

Ogcocephalus  nasutiis  (non  Cuvier  and  Valenciennes):  REID 
1954:73  [Cedar  Key,  Florida;  description:  ecology):  HER- 
ALD 1972:110  [color  photograph). 


Ogcocephalus  sp.:  SPRINGER  AND  WOODBURN  1960:91  [Tam- 
pa Bay,  Florida:  ecology]. 

Although  the  name  radiatus  Mitchill,  1818, 
has  often  been  used  for  this  form,  I  agree  with 
Longley  (Longley  and  Hildebrand  1941)  that  the 
usage  is  mistaken.  Longley  wrote  concerning 
Mitchill's  description  that  the  ".  .  .  facts  of  dis- 
tribution make  it  improbable  that  Mitchill's  Lo- 
phius  radiatus  is  the  same  [as  cubifrons 
Richardson]."  Unfortunately,  the  holotype  for 
radiatus  Mitchill  cannot  be  found.  The  fairly 
extensive  search  that  I  made  yielded  no  results, 
and  there  is  no  record  in  the  literature  or  else- 
where that  any  later  author  ever  saw  the  speci- 
men(s)  that  Mitchill  (1818)  described.  Therefore, 
we  have  only  the  original  description  of  radiatus 
on  which  to  rely,  but  it  is  impossible  to  tell  from 
it  what  Mitchill  had  in  hand.  All  but  one  of  the 
characters  in  the  original  description  are  generic 
or  familial  characters:  the  nature  of  the  modified 
scales  and  their  surface  sculpturing,  the  way  that 
scales  with  their  crystalline  tubercles  form  the 
rostrum,  the  location  of  the  rostral  cavity,  the 
"minute"  jaw  teeth,  the  "distinct"  tongue,  and 
the  size  and  placement  of  the  various  fins.  The 
one  character  given  which  is  helpful  in  discrim- 


260 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


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FIGURE  19.     Comparison  between  Ogcocephalus  cubifrons  and  O.  pantostictus  to  show  mouth  is  widest  in  O.  pantostictus 
relative  to  depth  of  head. 


inating  species  is  the  pectoral  fin  ray  count,  giv- 
en as  nine  or  ten.  But  these  numbers  are  too  low 
to  correspond  to  cubifrons,  for  which  the  range 
in  the  pectoral  fin  count  is  11  to  14.  However, 
Mitchill  (1818)  described  a  total  of  three  species 
of  ogcocephalids  in  his  paper  and  several  times 
gave  fin  counts  that  are  impossibly  low,  indicat- 
ing that  fin  ray  counts  cannot  be  used  with  con- 
fidence in  this  case.  The  locality  given  in  the 
original  description  of  radiatus  is  "the  depths 
of  the  Strait  of  Bahama."  Strait  of  Bahama  is  an 
old  name  for  the  Straits  of  Florida,  an  area  from 
which  could  be  taken  at  least  three  species  that 
fit  the  description;  these  are  rostellum,  nasutus, 
and  parvus,  the  latter  having  the  low  number  of 
pectoral  fin  rays  that  best  fits  the  description  if 
we  could  be  sure  that  it  were  accurate.  Of 
course,  cubifrons  also  is  taken  in  the  general 
area,  but  it  is  an  inshore  form  and  less  likely  to 
be  taken  in  "the  depths  of  the  Strait"  by  a  "drag 
net,"  if  we  interpret  that  to  mean  relatively  deep 
water  as  Longley  evidently  did  (Longley  and 
Hildebrand  1941).  Since  the  original  description 
of  Lophius  radiatus  Mitchill,  1818,  is  inadequate 


for  specific  assignment,  and  since  the  single  di- 
agnostic character  in  the  description  (the  pec- 
toral fin  ray  count)  fails  to  characterize  the 
species  at  hand,  I  hereby  reject  the  earlier  name 
in  favor  of  Lophius  (Malthe)  cubifrons  Richard- 
son, 1836. 

The  name  cubifrons  Richardson  is  represent- 
ed by  a  holotype,  BMNH  1855.9.19.848,  in  rea- 
sonable condition  except  for  fading,  and  the 
original  description  with  illustration  is  un- 
equivocally of  this  form,  but  there  is  a  question 
regarding  locality.  Jordan  and  Evermann  (1898) 
first  drew  attention  to  the  problem:  "Malthe 
cubifrons,  Richardson,  [is]  .  .  .  said  to  be  from 
Labrador  (Coll.  J.  J.  Audubon),  but  this  is  cer- 
tainly an  error;  Audubon  collected  also  in  Car- 
olina and  Florida."  Labrador  is  indeed  far  be- 
yond the  range  of  this  species,  but  "Carolina" 
and  Florida  are  both  likely  type-localities.  How- 
ever, I  am  unable  to  assign  the  holotype  to  any 
specific  locality,  or  even  general  locality,  within 
the  range  of  the  species. 

DIAGNOSIS  AND  COMPARISONS. — A  large 
species  of  Ogcocephalus  reaching  230  mm  SL 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


261 


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FIGURE  20.     Comparison  between  Ogcocephalus  rostellum  and  O.  cubifrons  to  show  that  depth  of  head  is  greatest  in  O. 
cubifrons  relative  to  length  of  disk  margin. 


(Table  1)  and  distinguished  by  the  combined  fea- 
tures of  a  bright  pattern  of  dark  spots  on  the 
pale  pectoral  fins,  a  small  mouth  (Fig.  6),  a  wide 
interorbital  distance,  and  a  thick,  massive  head 
on  which,  in  large  individuals,  is  perched  a 
small,  knoblike  rostrum,  because  the  rostrum  is 
characterized  by  ontogenetic  allometry:  very 
long  in  juveniles,  very  short  in  large  adults  (Fig. 
22). 

In  general  appearance,  O.  cubifrons  most 
closely  resembles  nasutus,  rostellum,  and  pan- 
tostictus,  especially  the  latter,  which  is  also 
large  and  heavy.  O.  cubifrons  is  distinguished 
from  pantostictus  by  its  smaller  mouth  (Fig.  19) 
and  lower  count  of  lateral-line  scales  in  the  sub- 
opercular  series  (usually  7-8  in  cubifrons,  mod- 
ally  9  in  pantostictus,  Table  6).  O.  cubifrons  is 
distinguished  from  both  pantostictus  and  nasu- 
tus by  the  lower  number  of  lateral-line  scales  in 
the  cheek  series  (modally  8  in  cubifrons,  9  in 
pantostictus  and  nasutus,  Table  5),  and  is  fur- 
ther distinguished  from  nasutus  by  having  a 
shorter  jaw  and  wider  interdrbital  (Figs.  14  and 
15)  and  in  having  allometric  growth  of  the  ros- 
trum (Fig.  22). 

Although  the  ranges  of  cubifrons  and  rostel- 
lum overlap  off  the  southeastern  United  States, 


the  two  evidently  seldom  co-occur,  cubifrons 
being  taken  in  shallower  water  than  rostellum. 
O.  rostellum  has  a  more  flattened  disk  as  ex- 
pressed by  the  head  height  (Fig.  20),  a  less  mas- 
sive head  as  expressed  by  the  rostrum  measure- 
ment (Fig.  9),  and  a  smaller  caudal  peduncle 
(Fig.  21);  it  is  a  smaller  species  than  cubifrons, 
reaching  only  160  mm  SL. 

Ogcocephalus  parvus  and  corniger  occur  off 
both  coasts  of  Florida  and  off  the  Carolinas  and 
Yucatan,  as  does  cubifrons,  but  parvus  and  cor- 
niger are  taken  in  deeper  water  than  cubifrons. 
Both  parvus  and  corniger  have  fewer  pectoral 
fin  rays  than  cubifrons  (usually  12-13  in  cubi- 
frons, 10-11  in  parvus  and  corniger,  Table  3) 
and  differing  color  patterns;  parvus,  a  small 
species  reaching  only  85  mm  SL,  has  blotches 
of  varying  tones  as  the  color  pattern  on  the  dor- 
sal surface  of  the  body,  but  it  lacks  the  promi- 
nent clusters  of  dark  circles  seen  on  the  body 
and  pectoral  fins  in  cubifrons.  O.  corniger  dif- 
fers from  cubifrons  in  having  close-set  pale 
spots  on  a  dark  background  evenly  covering  the 
entire  dorsal  surface  of  the  body. 

Ogcocephalus  declivirostris  is  restricted  to 
the  northwestern  Gulf  of  Mexico  and  is  evi- 
dently allopatric  with  cubifrons.  O.  decliviros- 


262 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


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FIGURE  21.     Comparison  between  Ogcocephalus  rostellum  and  O.  cubifrons  to  show  that  depth  of  caudal  peduncle  is 
greatest  in  O.  cubifrons  relative  to  length  of  disk  margin. 


tris  is  distinguished,  in  any  case,  in  having  fewer 
pectoral  fin  rays  (modally  11  in  dedivirostris, 
usually  12-13  in  cubifrons,  Table  3);  dediviros- 
tris lacks  the  fleshy  ventral  pads  on  the  distal 
tips  of  the  pectoral  fin  rays  as  well  as  the  spotted 
pattern  of  the  pectoral  fins  so  prominent  in  cub- 
ifrons (although  occasionally  dedivirostris  has 
a  few  very  pale  spots  on  the  pectorals).  Finally, 
dedivirostris  differs  from  cubifrons  in  having  its 
rostrum  jutting  horizontally  or  downwards  in- 
stead of  upwards  as  in  cubifrons. 

DESCRIPTION. — Counts  and  measurements 
from  95  specimens  29.9  to  224.0  mm  SL  (Table 
1). 

Counts.  Counts  given  in  Tables  2-6.  Two 
counts  for  cubifrons  lie  in  the  middle  of  the 
range  for  the  genus;  pectoral  fin  ray  count  is 
generally  12-13  in  cubifrons,  which  is  the  mid- 
dle of  the  generic  range  of  10-15,  and  the  sub- 
opercular  lateral-line  scale  count  in  cubifrons  is 
usually  7-8,  which  is  the  middle  of  the  generic 
range  (excluding  extremes)  of  5-10.  The  lateral- 
line  scale  count,  although  showing  wide  varia- 
tion, is  also  near  the  middle  of  the  range  for  the 


genus  (Table  4).  However,  cubifrons  agrees 
with  pantostictus,  rostellum,  and  dedivirostris 
in  having  a  high  modal  vertebral  count  of  20, 
whereas  the  more  usual  count  for  the  genus 
is  19. 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses. 

Length  of  disk  margin  intermediate  within 
range  of  variation  for  the  genus,  2.3(2.0-2.5)  in 
SL.  Width  of  tail  also  intermediate,  varying  from 
thin  in  smallest  specimens  to  thick  and  broad  in 
large  specimens,  1.8(1.5-2.4)  in  length  of  disk 
margin;  caudal  peduncle  thick  and  deep,  its 
depth  2.5(2.2-2.9)  in  head  depth.  Rostrum  long, 
conical,  and  upturned  in  small  specimens,  be- 
coming relatively  small  and  knoblike  in  large 
ones  (Fig.  22);  length  of  rostrum  4.3(1.8-6.1)  in 
length  of  disk  margin.  Aperture  of  illicial  cavity 
subtriangular  in  small  specimens,  much  higher 
than  wide,  becoming  merely  oval  or  round  in 
large  ones.  Cranium  rising  steeply  above  the 
disk,  its  contours  becoming  massive  and  boxlike 
in  adults;  head  depth  2.0(1.7-2.2)  in  length  of 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


263 


43  mm 


71  mm 


135  mm 


304  mm 


FIGURE  22.     Heads  of  four  examples  of  Ogcocephalus  cub- 
ifrons  showing  allometric  growth  of  rostrum. 


disk  margin.  Width  of  cranium  3.1(2.6-3.6)  in 
length  of  disk  margin.  The  following  in  width  of 
cranium:  eye  1.6(1.4-1.9),  lateral  ethmoid  width 
1.8(1.5-2.0).  Interorbital  very  wide,  slightly 
concave,  1.9(1.5-2.2)  in  width  of  cranium, 
3.0(2.3-3.4)  in  head  depth.  Mouth  small,  its 
width  1.9(1.6-2.2)  in  head  depth,  3.8(3.2-4.4)  in 
length  of  disk  margin;  jaw  2.8(2.5-3.2)  in  head 
depth.  Upper  lip  fleshy,  of  an  even  width  for  its 
entire  length;  lower  lip  fleshy,  usually  thickened 
medially  although  no  median  lobe  is  developed. 
Dorsal  fin  3.4(2.7-4.8)  in  length  of  disk  margin; 
anal  fin  2.6(2.2-3.1)  in  length  of  disk  margin. 

Integument.  Dermal  cirri  generally  well  de- 
veloped on  lateral  sides  of  tail,  less  so  on  disk 
margin;  cirri  also  frequently  scattered  thickly  on 
dorsal  surface  of  body,  particularly  in  half- 
grown  specimens.  Large  bucklers  prominent, 
arranged  as  in  generic  description.  Tubercles 
relatively  sparse  on  fins,  occurring  at  bases  of 
dorsal  and  pectoral  fins  with  only  a  few  scattered 
along  fin  rays.  Anal  and  ventrals  with  tubercles 
at  bases  and  running  out  on  fin  rays  for  a  third 
to  a  half  their  length.  Caudal  as  in  generic  de- 
scription. Skin  covering  eyeballs  bearing  tiny 
tubercles  with  rim  of  cornea  beaded  by  a  row  of 
small  bucklers.  Pectoral  membranes  thick, 
opaque,  the  ventral  surface  of  ray  tips  with  well- 
developed  fleshy  pads. 

Color  in  preservative.  Ground  color  of  dorsal 
surface  of  body  light  tan  to  dark  brown  or  gray 
with  black  spots  sprinkled  on  face,  on  skin  cov- 
ering eyeballs,  on  shoulders  and  lateral  sides  of 
tail,  in  the  axillae,  on  pectoral  fins,  and  usually 
on  lateral  margins  of  disk.  Spotted  pattern  on 
shoulders  does  not  form  a  reticulum  as  it  does 


in  O.  vespertilio.  Shoulder  spots  often  with  pale 
centers  in  small  specimens.  Ground  color  of 
pectoral  fins  and  sides  of  tail  usually  pale  with 
dark  spots  so  close  together  that  the  background 
appears  as  a  reticulum;  even  in  specimens  so 
dark  as  to  obscure  most  of  the  spotted  pattern 
of  body,  the  pattern  on  pectoral  fins  remains 
prominent.  Pectoral  fins  with  a  broad  dark 
brown  or  black  border  in  medium-sized  and 
small  specimens.  Lips  pale;  rostrum  with  a  dif- 
fuse black  subterminal  ring;  iris  black,  or  golden 
with  black  spots,  in  which  case  color  pattern  of 
face  extends  over  eyes  to  margin  of  pupil.  Ven- 
tral surface  of  body  uniformly  pale  or  creamy. 
Ventral  and  anal  fins  the  same  ground  color  as 
ventral  surface  of  body,  ventrals  often  with  a 
pale  patch  distally  in  dark  specimens,  anal  fin 
sometimes  dark  or  at  least  tipped  with  brown  or 
black  in  small  specimens.  Ventral  pads  of  pec- 
toral fin  rays  white  or  pale.  Dorsal  fin  spotted  or 
sometimes  with  merely  vague  blotches.  Caudal 
fin  as  in  generic  description  but  generally  with 
spotted  pattern  of  tail  carried  back  and  super- 
imposed on  caudal's  basic  pattern. 

Color  in  life.  I  have  not  seen  this  species  alive. 
The  following  is  a  description  by  Longley 
(Longley  and  Hildebrand  1941),  "The  young  are 
dark  gray  on  the  under  side;  older  fish  coppery 
red,  with  much  individual  variation  in  color  on 
the  dorsal  side,  usually  lighter  or  darker  brown, 
variegated  with  reddish  orange  or  yellow-orange 
patches,  disappearing  in  preserved  specimens; 
the  chief  patches  behind  the  eyes,  the  two  some- 
times meeting  across  the  nape  to  form  a  great 
U-shaped  mark  open  to  the  rear;  usually  an  ad- 
ditional one  above  and  behind  the  branchial  ap- 
erture; and  one  about  base  of  soft  dorsal  fin. 
Even  at  a  standard  length  of  50  mm.  there  is 
some,  and  in  fish  of  medium  and  large  size  much 
spotting  with  light-bordered  dark  spots  in  a  long, 
narrow  tract  from  eye  to  gill  opening.  Here  the 
spots  usually  are  less  crowded  than  elsewhere, 
and  more  nearly  circular.  On  the  sides  of  the 
head,  margin  of  the  disk,  and  sides  of  the  trunk, 
the  pattern  is  less  open,  being  principally  a  net- 
work of  light  lines  on  the  dark  ground.  Small 
fish  have  pectorals  and  caudal  yellow  proximally 
and  dark  brown  distally,  with  a  netted  pattern 
appearing  first  at  the  base  of  these  fins  and  ex- 
tending gradually  to  their  tips,  the  yellow  ad- 
vancing before  it  at  the  expense  of  the  darker 
brown,  which  gives  way  to  the  lighter  color  on 
which  the  marbling  spreads." 


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A  photograph  in  Walls  (1975:263)  shows  a 
specimen  (his  O.  radiatus)  with  margins  of  pec- 
torals bright  yellow;  the  size  of  the  fish  is  not 
given,  but  the  small  relative  size  of  the  rostrum 
indicates  that  the  specimen  is  probably  a  large 
adult.  A  smaller  specimen  is  shown  in  color  in 
Herald  (1972).  In  this  specimen  the  yellow  color 
lies  just  proximal  to  the  dark  brown  fin  margins 
about  as  Longley  described  (Longley  and  Hil- 
debrand  1941).  Both  photographs  show  the 
"netted"  pattern  on  the  base  of  the  pectorals. 

DISTRIBUTION  (Fig.  17). — Northern  Bahamas, 
coast  of  the  southeastern  United  States  from  the 
vicinity  of  Cape  Lookout,  North  Carolina, 
around  the  Florida  peninsula  to  Pensacola,  Flor- 
ida, and  south  to  Campeche  Banks,  Yucatan. 
Bathymetric  range:  shore  to  68  m. 

MATERIAL  EXAMINED.— BMNH  1855.9.19.848,  the  holo- 
type,  said  to  be  from  Labrador,  but  this  undoubtedly  an  error 
(see  p.  260).  For  following  material,  numbers  in  parentheses 
are  numbers  of  specimens.  For  data  of  OREGON  stations  (  = 
O),  refer  to  Springer  and  Bullis  (1956). 

United  States:  North  Carolina:  UF  24912  (1),  near  Cape 
Lookout  Bight,  Carteret  Co.  South  Carolina:  UF  24903  (1), 
31°58'N,  79°43'W,  about  77.3  km  SE  Edisto  Beach,  Beaufort 
Co.  Georgia:  UF  25931  (1),  off  Brunswick.  Florida:  Dade 
County:  FMNH  46128  (1),  inshore  reef  between  Miami  and 
Angelfish  Creek;  FMNH  46854  (1),  Triumph  Reef  off  Elliot 
Key;  USNM  89710-89711  (2),  Biscayne  Bay  near  Cocoanut 
Grove,  Miami.  Monroe  County:  ANSP  90667  (1),  E  end 
Grassy  Key,  Florida  Bay;  CAS  18585  (1),  near  Key  West; 
FMNH  7020  (1),  Key  West;  FMNH  21645-21647  (3),  near 
Marathon;  FMNH  64123  (1),  O-1004;  FSBC  342  (4),  FSBC 
381  (1),  FSBC  410  (1)  all  from  24041'-24°52'N,  82<W- 
82°35'W;  FSBC  626  (1)  and  FSBC  762  (1),  both  from 
24°43'10"-24°52'00"N,  82°00'-82°35'W;  UF  4071  (1),  Marque- 
sas Keys;  UF  7022  (1),  Big  Pine  Key;  UMML  1201  (10), 
24°45'-24°50'N,  82°10'-82°30'W;  UMML  4786  (1),  5.3  km  sw 
of  Greyhound  Key;  USNM  57243  (1),  Matecumbe  Key: 
USNM  84035  (1),  24°36'N,  81°48'W;  USNM  117003  (7),  Tor- 
tugas;  USNM  1 17006  (3),  channel  w  of  White  Shoal,  Tortugas; 
USNM  117170  (2),  Tortugas;  USNM  126488  (2),  Key  West. 
Collier  County:  FSBC  1157  (1),  Everglades.  Lee  County: 
CAS-SU  5841  (1),  Captiva  Pass;  CAS-SU  36404  (6),  Sanibel 
I.;  FMNH  510,  512,  and  513  (3),  Captiva  Pass:  FMNH  38584- 
38586  (3)  and  FMNH  38587  (1),  Boca  Grande;  USNM  76025 
(1),  Boca  Grande  Pass:  USNM  86159(1),  Boca  Grande.  Hills- 
borough  County:  USNM  143160  (2),  Port  Tampa.  Pinellas 
County:  FSBC  61  (1),  Blind  Pass,  St.  Petersburg  Beach:  FSBC 
878  (1),  John's  Pass,  Madeira  Beach:  FSBC  1369  (1),  due  w 
of  Don-ce-Sar;  USNM  158543  (1),  Bay  at  Clearwater.  Levy 
County:  the  following  all  from  Cedar  Key:  ANSP  90668  (2); 
CAS-SU  99  (5):  UF  229  (1):  UF  753  (1):  UF  916  (1);  UF  919 
(1);  UF  1522  (1):  UF  1612  (1):  UF  3169  (1):  UF  3374  (1):  UF 
7832  (1);  UF  (1),  R-I-650-4:  UF  (1),  R-l-651-3:  UF  (2),  R-I- 
2351-1;  UF(l),R-II-551-6:  UF(1),  R-1I-2051-4:  UF(1),  R-III- 
451-1:  UF(1),  R-III-451-4;  UF(2),  R-IV-551-1:  UF(1),  R-IV- 
551-7:  UF  (2),  R-V1-650-3:  UF  (4).  R-VI-750-2;  UF  (4),  R-VI- 
1750-4;  UF(1),  R-VII-150-5;  UF  (4),  R-VII-150-6;  UF  (6),  R- 
VIII-2650-1;  UF  (4),  R- VII 1-2650-1;  UF  (2),  R-VIII-2650-2; 


UF  (1),  R-IX-1250-1;  UF  (1),  R-IX-1250-1;  UF  (2),  R-X-850- 
7;  UF  (1),  R-X-2250-5-5;  UF  (1),  R-XI-750-6;  UF  (1),  R-XI- 
750-7;  UF  (1),  R-XII-1050-1;  UF  (2);  R-XII-1050-2;  USNM 
88499  (1);  USNM  164928  (1).  Franklin  County:  CAS  23914 
(1),  Apalachicola  Bay.  Okaloosa  County:  UF  5586  (1),  UF 
5587  (1),  UF  5596  (1),  all  from  Choctawatchee  Bay  at  Destin; 
UF  2730(1),  Pensacola. 

Florida  Straits:  UF  28649  (1),  SILVER  BAY  sta.  144,  27°44'N, 
82°46'W,  7.3  m. 

Bahamas:  ANSP  103627  (1),  GEORGE  M.  BOWERS  sta.  659, 
w  side  Eleuthra  I.;  ANSP  (3),  Chaplin  Program  sta.  513B, 
Hatchet  Bay,  Eleuthra  I.;  ANSP  (2),  Chaplin  Program  sta. 
526,  Hatchet  Bay,  4.8  km  offshore,  Eleuthra  I.:  UMML  4719 
(1),  Exuma  Sound  adjacent  to  Eleuthra  I.,  7.3  m. 

Mexico:  Campeche  Banks:  FMNH  46745  (5),  O-436;  FMNH 
64107  (8),  O-713-716;  FMNH  64134  (3),  O-710;  UF  28647  (1), 
SILVER  BAY  sta.  362,  20°00'N,  91°43'W,  44  m;  UF  28648  (1), 
O-417.  Yucatan:  USNM  148497  (1). 

Undetermined  material.  Seven  specimens  from  the  Bahama 
Islands  are  intermediate  in  color  pattern  between  cubifrons 
and  nasutus,  and  also  cannot  be  diagnosed  with  certainty  on 
the  basis  of  morphometric  characters.  These  specimens  are: 
ANSP  72335  (1),  Hog  I.;  ANSP  90670-90671  (2),  s  shore  of 
Hog  I.;  and  FMNH  64139  (4),  Grand  Bahama  I. 

Ogcocephalus  pantostictus  new  species 

(Figure  23) 

Ogcocephalus  vespertilio  (non  Linnaeus):  WOODS  1942:192 
[off  Corpus  Christi,  Texas]. 

Ogcocephalus  cubifrons  (non  Richardson):  HILDEBRAND 
1954:321  [off  Texas]. 

Ogcocephalus  radiatus:  ANONYMOUS  1976:131  and  427  [Mex- 
ican coast  of  Gulf  of  Mexico;  color  photograph;  key]; 
HOESE  AND  MOORE  1977:144,  fig.  101  [NW  Gulf  of  Mexico; 
key;  partial  description;  range]. 

DIAGNOSIS  AND  COMPARISONS. — A  very  large 
Ogcocephalus  (Table  1)  with  the  entire  dorsal 
surface  of  the  body  covered  with  a  polkadotlike 
pattern  of  dark  spots  (sometimes  dim  in  museum 
specimens).  Resembling  cubifrons  in  many  fea- 
tures, it  has  allometric  growth  of  the  rostrum: 
a  relatively  long  rostrum  in  small  specimens,  but 
short  and  sometimes  merely  knoblike  in  large 
specimens.  O.  pantostictus  is  distinguished 
from  cubifrons  by  its  wider  mouth  (Fig.  19;  also 
expressed  by  the  jaw  measurement,  Fig.  14)  and 
in  having  higher  modal  counts  of  lateral-line 
scales  in  the  cheek  series  (9  in  pantostictus,  8 
in  cubifrons,  Table  5)  and  in  the  subopercular 
series  (9  in  pantostictus,  8  in  cubifrons,  Table 
6). 

With  its  range  restricted  to  the  northern  and 
western  Gulf  of  Mexico,  pantostictus  is  largely 
sympatric  with  declivirostris,  but  the  two  are 
distinguished  by  the  pectoral  fin  ray  count  (usu- 
ally 12-13  in  pantostictus  and  11  in  decliviros- 
tris, Table  3)  and  the  lateral-line  scale  count  in 
the  subopercular  series  (modally  9  in  pantostic- 
tus, 1  in  declivirostris,  Table  6),  O.  pantostictus 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


265 


FIGURE  23.  Ogcocephalus  pantostictus  new  species,  all  from  Gulf  of  Mexico.  (A)  and  (D)  Paratype  77.0  mm  SL,  CAS-SU 
40559,  Galveston,  Texas.  (B)  Paratype  96.0  mm  SL,  FMNH  64111,  off  Mobile  Bay,  Alabama.  (C)  and  (E)  Holotype  226.0  mm 
SL,  FMNH  50765,  off  Pensacola,  Florida.  (F)  Paratype  187.0  mm  SL,  CAS-SU  40559,  showing  variation  of  rostrum  resembling 
O.  cubifrons.  Compare  with  (G),  O.  cubifrons  (same  specimen  as  in  Fig.  18C). 


also  has  the  ventral  surfaces  of  the  pectoral  fin 
rays  thickened  with  fleshy  pads  which  are  not 
present  in  declivirostris. 

The  species  corniger  and  parvus  have  ranges 
slightly  overlapping  that  of  pantostictus  in  the 
northern  Gulf  of  Mexico,  but  both  are  separable 
from  pantostictus  by  their  lower  pectoral  fin  ray 
counts  (usually  12-13  in  pantostictus  and  10-1 1 
in  corniger  and  parvus,  Table  3).  O.  parvus  is 
a  far  smaller  species  than  pantostictus,  reaching 
only  about  85  mm  SL.  O.  corniger  has  a  much 
longer  rostrum  than  does  pantostictus  (Fig.  9) 
and  has  a  distinctive  color  pattern  of  pale  spots 
on  a  darker  background  distributed  evenly  over 
the  dorsal  surface  of  the  body  in  contrast  to  the 
darkish  circles  on  a  lighter  background  found  in 
pantostictus. 


DESCRIPTION. — Counts  and  measurements 
from  65  specimens,  38.6  to  261.0  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6;  in  the 
following  description,  counts  for  the  holotype 
given  in  brackets.  Pectoral  fin  ray  counts  in  pan- 
tostictus in  middle  of  range  for  genus,  12-13  in 
a  range  of  10-15  [12,12].  Subopercular  lateral- 
line  scale  count,  modally  9  [10,10]  in  pantostic- 
tus, high  relative  to  range  of  5-10  (excluding  ex- 
tremes) for  genus;  only  two  other  species  have 
as  high  a  count:  vespertilio  usually  has  9  scales 
in  subopercular  series,  notatus  usually  10.  The 
count  for  cheek  lateral-line  scales  nears  high  end 
of  range,  too,  with  count  in  pantostictus  usually 
8  or  9  [9,10];  in  this  character  pantostictus 
agrees  with  nasutus,  and  only  vespertilio  and 


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notatus  have  higher  counts  in  cheek  series.  Lat- 
eral-line scale  count  [35,36]  varies  widely,  but 
mean  is  highest  for  genus.  O.  pantostictus  re- 
sembles cubifrons,  rostellum,  and  declivirostris 
in  vertebral  count  of  19-21,  modally  20,  in  con- 
trast to  remaining  eight  species  which  nearly  al- 
ways have  19,  occasionally  18,  vertebrae.  [D.4; 
A.4.] 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses and  the  proportion  for  the  holotype  in 
brackets. 

Length  of  disk  margin  intermediate  within 
range  of  variation  for  the  genus,  2.3(2.1- 
2.4)[2.4]  in  SL.  Tail  moderately  thick  and  broad 
in  large  specimens,  relatively  narrow  in  small 
ones,  its  width  1.8(1.5-2.0)[1.7]  in  length  of  disk 
margin;  caudal  peduncle  thick  and  deep,  its 
depth  2.6(2. 3-2.9)[2.5]  in  depth  of  head.  Onto- 
genetic  allometry  in  growth  of  rostrum  resem- 
bles that  in  cubifrons ;  however,  I  have  not  seen 
specimens  of  pantostictus  less  than  38.6  mm  in 
standard  length  and  therefore  do  not  know 
whether  or  not  the  remarkably  long  rostrum 
seen  in  very  small  cubifrons  (30  mm  SL)  is  also 
characteristic  of  small  pantostictus.  Rostrum 
cone-shaped  in  small  examples,  becoming  rela- 
tively small  and  finger-shaped  in  large  ones. 
Length  of  rostrum  4.3(2.6-6.3)[5.0]  in  length  of 
disk  margin.  Aperture  of  illicial  cavity  subtrian- 
gular  in  small  specimens,  higher  than  wide,  be- 
coming oval  in  adults.  Cranium  rising  steeply 
above  disk,  becoming  heavy  and  massive  in 
large  specimens;  head  depth  2.0(1.8-2.1)[2.1]  in 
length  of  disk  margin.  Width  of  cranium  3.2(2.9- 
3.7)[3.4]  in  length  of  disk  margin.  The  following 
in  width  of  cranium:  eye  1.6(1.4-1.8)[1.8],  lat- 
eral ethmoid  width  1. 6(1.4-1. 8)[1.6].  Interorbital 
slightly  concave,  wide,  2.0(1.8-2.5)[1.9]  in  width 
of  cranium,  3.3(2.9-4.1)[2.9]  in  head  depth. 
Mouth  moderate,  its  width  1.6(1.4-1.8)[1.6]  in 
head  depth,  3.1(2.8-3.6)[3.3]  in  disk  margin;  jaw 
2.3(2.1-2.7)[2.5]  in  head  depth.  Upper! lip  fairly 
fleshy,  of  an  even  width  along  its  entire  length; 
lower  lip  fleshy,  usually  thickened  medially  al- 
though no  median  lobe  is  developed.  Dorsal  fin 
3.7(2.9^.8)[4.8]  in  length  of  disk  margin;  anal 
fin  2.7(2.3-3.2)[3.2]  in  length  of  disk  margin. 

Integument.  Dermal  cirri  often  present  on 
disk  margin  and  lateral  sides  of  tail.  Large  buck- 
lers prominent,  arranged  as  in  generic  descrip- 
tion. Tubercles  relatively  sparse  on  fins,  occur- 
ring at  very  base  in  dorsal  and  anal  fins  and  on 
about  proximal  third  of  anal  and  ventrals.  Cau- 


dal as  in  generic  description.  Skin  covering  eye- 
balls bearing  tubercles  with  rim  around  cornea 
usually  beaded  by  a  row  of  small  bucklers.  Pec- 
toral membranes  thick,  opaque;  ventral  surfaces 
of  ray  tips  with  well-developed  fleshy  pads. 

Color  in  preservative.  Dorsally  the  ground 
color  of  body  gray  or  tan  sprinkled  fairly  evenly 
with  round  dark  spots  over  entire  tail  and  disk, 
including  face  and  skin  covering  eyeballs.  Mark- 
ings on  shoulders,  sides  of  tail,  and  in  axillae 
generally  darkest  and  showing  greatest  contrast 
with  background  color.  Ventral  surface  uniform- 
ly pale,  usually  paler  than  ground  color  of  dorsal 
body  surface.  Lips  pale;  rostrum  with  a  diffuse 
dark  subterminal  ring;  iris  golden  with  black 
spots,  or  entirely  black. 

Pectorals  with  same  spotted  pattern  as  dorsal 
surface  of  body.  In  small  or  medium-sized  spec- 
imens, spotted  pattern  on  pectorals  grades  into 
a  dark  border.  In  many  individuals  spots  close 
together  so  that  background  appears  as  a  light- 
colored  reticulum.  Ventrals  and  anal  same  color 
as  ventral  surface  of  body;  anal  sometimes 
dusky  near  tip  in  specimens  under  about  100  mm 
SL.  Dorsal  fin  usually  spotted,  especially  when 
spotted  pattern  of  dorsal  body  surface  relatively 
intense. 

Color  in  life.  Unknown. 

DISTRIBUTION  (Fig.  17). — Coasts  of  northern 
and  western  Gulf  of  Mexico  from  approximately 
longitude  87°W  to  latitude  25°N.  Bathymetric 
range:  from  limited  data  available,  about  9- 
31  m. 

NAME. — The  name  pantostictus  means  all 
spotted,  in  reference  to  the  distinctive  over-all 
spotted  color  pattern. 

MATERIAL  EXAMINED. — HOLOTYPE:  FMNH  50765,  Santa 
Rosa  Sound,  Pensacola,  Florida;  a  male  226.0  mm  SL. 

PARATYPES.  Numbers  in  parentheses  are  numbers  of  speci- 
mens. United  States:  Florida:  ANSP  71329  (1),  Pensacola; 
MCZ  45080  (1),  OREGON  sta.  1648,  30°11'N,  86°55'W,  31  m. 
Alabama:  FMNH  64111  (1),  OREGON  sta.  882,  30°10'N, 
87°50'W,  10  m.  Mississippi:  CAS  23934  (1),  off  Horn  I.; 
FMNH  21636  (1),  Horn  I.;  FMNH  46734  (1),  outside  Petit 
Bois  I.;  GCRL  562  (1),  outside  Petit  Bois  I.;  GCRL  563  (1), 
sw  of  Ship  I.;  GCRL  564  (1),  off  Horn  I.;  GCRL  565  (1),  off 
w  end  of  Horn  I.;  UF  uncat.  (1),  SILVER  BAY  sta.  167, 
30°12'N,  88°40'W,  7.3  m.  Louisiana:  FMNH  50997  (2);  GCRL 
566  (1),  s  of  Grand  Isle,  17  m;  USNM  188755  (1),  PELICAN 
sta.  66-7,  28°59'N,  91°48'W.  Texas:  CAS  23932  (1);  USNM 
94443  (1);  USNM  118610(9).  Vicinity  ofGalveston:  CAS-SU 
40559  (10);  USNM  118603  (1);  USNM  118604  (1);  USNM 
118606  (1);  USNM  120071  (1);  USNM  176982  (1);  USNM 
188803  (2),  PELICAN  sta.  56-5,  29°10.5'N,  94°50.5'W;  USNM 
188805  (1),  PELICAN  sta.  62-2,  29°19'N,  94°31.5'W.  Vicinity  of 
Freeport:  MCZ  45079  (1),  USNM  1 17804  (14),  USNM  1 18605 
(1),  USNM  118611  (1).  Southern  Texas:  USNM  120070  (1), 


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267 


FIGURE  24.  (A)  and  (B)  Ogcocephalus  rostellum  new  species,  holotype,  133.0  mm  SL,  USNM  188809,  off  Atlantic  coast 
of  Florida,  29°10'N,  80°19'W.  (C)  and  (D)  Ogcocephalus  declivirostris  new  species,  holotype,  129.5  mm  SL,  FMNH  64136, 
Texas  coast  of  Gulf  of  Mexico,  28°02'N,  %°03'W.  (E)  Paratype  of  O.  declivirostris,  107.5  mm  SL,  FMNH  61577,  Texas  coast 
of  Gulf  of  Mexico,  28°02'N,  96°04'W,  showing  pattern  of  dim  spots  sometimes  seen  on  dorsal  surface  of  body. 


Aransas  Pass;  FMNH  37728-37729  (2),  Port  Aransas; 
FMNH  40298  (1),  within  80  km  of  Corpus  Christi;  ANSP 
90675  (1),  Brownsville  Ship  Channel,  Pt.  Isabel;  USNM 
188800(1),  Pt.  Isabel. 

Mexico:  USNM  188775  (1)  and  USNM  188783  (1),  both  from 
off  Huts  Bayon,  approximately  24°30'N,  6-18  m. 

Material  without  exact  localities:  USNM  188756  (1),  PELI- 
CAN sta.  37;  USNM  188758  (1),  PELICAN  sta.  104-2;  USNM 
188769  (1),  MC  9;  USNM  188798  (1),  8.1  km  offshore. 

Ogcocephalus  rostellum  new  species 

(Figure  244 -B) 

Malthaea  nasuta  (non  Cuvier  and  Valenciennes):  DEKAY 
1842:166,  pi.  28,  fig.  89  [synonymy;  description;  range, 
compiled]. 

DIAGNOSIS  AND  COMPARISONS. — A  moderate- 


sized  Ogcocephalus  with  a  very  abbreviated  ros- 
trum resembling  the  shelflike  condition  seen  in 
notatus.  (In  the  smallest  specimens  of  both  ros- 
tellum and  notatus,  40  mm  SL  or  less,  a  short 
median  horn  is  developed  on  the  rostrum.)  O. 
rostellum  is  distinguished  from  notatus  by  its 
narrower  mouth  (Fig.  6)  and  lower  number  of 
scales  in  the  cheek  lateral-line  series  (modal 
number  8  in  rostellum,  10  in  notatus,  Table  5) 
and  in  the  premaxillary  series  (3  in  rostellum,  4 
in  notatus). 

In  general  appearance  and  color  pattern,  ros- 
tellum most  closely  resembles  cubifrons;  the 
ranges  of  the  two  species  overlap  along  the  At- 


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lantic  coast  of  the  southeastern  United  States, 
but  rostellum  is  ordinarily  taken  from  deeper 
water  than  cubifrons.  The  two  are  distinguished 
by  some  differences  in  body  proportions:  ros- 
tellum has  a  relatively  more  flattened  disk  as 
expressed  by  the  head  height  measurement  (Fig. 
20)  and  a  less  massive  head  as  expressed  by  the 
rostrum  measurement  (Fig.  10);  also  it  has  a  nar- 
rower caudal  peduncle  than  cubifrons  (Fig.  21). 
O.  rostellum  is  a  smaller  species  than  cubifrons, 
reaching  160.0  mm  in  standard  length  compared 
to  230.0  mm  in  cubifrons.  Finally,  the  color  pat- 
tern of  the  pectoral  fin  helps  discriminate  be- 
tween rostellum  and  cubifrons;  often  there  are 
no  dark  spots  on  the  pectoral  fins  in  rostellum 
compared  to  the  bright  pattern  of  dark  spots  on 
a  light  background  covering  the  pectoral  fins  in 
cubifrons,  but  when  spots  do  occur  on  the  pec- 
torals in  rostellum,  they  are  generally  dim  and 
also  less  numerous  than  in  cubifrons  and  re- 
stricted to  the  basal  portions  of  the  fins. 

Ogcocephalus  rostellum  has  its  range  adjacent 
to  that  of  nasutus  but  is  a  smaller  species  than 
nasutus  (Table  1)  and  lacks  the  long  rostrum 
seen  in  nasutus  (Fig.  10);  rostellum  differs  fur- 
ther from  nasutus  in  having  a  shorter  jaw  (Fig. 
16). 

The  range  of  rostellum  overlaps  the  ranges  of 
parvus  and  corniger  \  all  occur  at  moderate 
depths  on  the  continental  shelf  off  the  south- 
eastern United  States.  O.  rostellum  may  be  sep- 
arated from  parvus  and  corniger  by  the  pectoral 
fin  ray  count  (usually  13  in  rostellum,  usually 
10-11  in  parvus  and  corniger,  Table  3).  O.  ros- 
tellum is  further  distinguished  from  corniger  in 
lacking  a  long  rostrum  (Fig.  9),  and  is  further 
distinguished  from  parvus  in  its  flatter  disk  with 
less  elevated  cranium  (head  depth  4.3-5.4  into 
SL  in  rostellum,  3.1-4.2  in  parvus). 

DESCRIPTION. — Counts  and  measurements 
from  69  specimens,  33.3  to  154.2  mm  SL  (Table 
1). 

Counts.  Counts  given  in  Tables  2-6;  in  the 
following  description,  counts  for  holotype  given 
in  brackets.  Pectoral  fin  ray  count  usually  13  [13, 
13],  range  11-15,  about  intermediate  within 
range  of  variation  for  genus.  Count  of  lateral- 
line  scales  in  subopercular  series  moderately 
low  for  genus,  usually  6  or  7  [5,6],  range  5-9. 
Count  of  cheek  series  of  lateral-line  scales  8 
[8,8],  the  most  common  count  for  the  genus; 
range  7-9.  Lateral-line  scale  count,  21-32  [26, 


25].  Mean  for  each  of  the  counts  of  series  of 
lateral-line  scales  in  rostellum  lower  than  in  the 
closely  similar  cubifrons.  O.  rostellum  agrees 
with  cubifrons,  pantostictus,  and  declivirostris 
in  having  a  modal  count  of  20  vertebrae,  com- 
pared to  other  species  in  genus  in  which  modal 
count  is  19.  [D.  4;  A.  4.] 

Proportions.  Proportions  expressed  as  ratios 
given  as  the  mean  followed  by  the  range  in  pa- 
rentheses and  the  proportion  for  the  holotype  in 
brackets. 

Length  of  disk  margin  intermediate  relative  to 
range  of  variation  for  genus,  2.2(2.0-2.4)[2.4]  in 
SL.  Tail  moderate  in  width,  1.8(1.6-2.1)[1.6]  in 
length  of  disk  margin;  caudal  peduncle  slender, 
2.7(2.4-2.9)[2.7]  in  head  depth.  Rostrum  pro- 
duced into  a  short  cone  in  small  specimens,  be- 
coming more  shelflike  in  large  specimens  with 
the  median  horn  extremely  abbreviated,  rostrum 
6.1(3.7-8.0)[7.1]  in  length  of  disk  margin.  Ap- 
erture of  illicial  cavity  almost  diamond-shaped. 
Cranium  well  demarcated  from  disk,  but  head 
less  deep  than  in  other  species  in  the  genus,  head 
depth  2.2(2.0-2.4)[2.2]  in  length  of  disk  margin. 
Cranium  relatively  narrow,  its  width  3.5(2.7- 
3.9)[3.6]  in  length  of  disk  margin  (one  specimen 
had  this  ratio  4.4).  The  following  in  width  of 
cranium:  eye  1.5(1.3-1.7)[1.5],  lateral  ethmoid 
width  1.6(1. 5- 1.8) [1.8].  Interorbital  space  slight- 
ly concave,  its  width  moderate,  2.2(1.9-2.7)[2.4] 
in  width  of  cranium,  3.4(2.8-4.1)[3.9]  in  head 
depth.  Mouth  small,  its  width  1.7(1.5-2.0)[1.8] 
in  head  depth,  3.8(3. 1-4.4)[4.0]  in  length  of  disk 
margin;  jaw  2.6(2.3-2.8)[2.7]  in  head  depth.  Up- 
per lip  fleshy,  of  an  even  width  for  its  entire 
length;  lower  lip  fleshy,  very  thickened  medial- 
ly. Dorsal  fin  4.3(3.2-4.9)[4.5]  in  length  of  disk 
margin;  anal  fin  3.0(2.5-3.5)[3.4]  in  length  of 
disk  margin. 

Integument.  Dermal  cirri  sometimes  present 
but  never  conspicuous.  Large  bucklers  promi- 
nent, arranged  as  in  generic  description.  In  more 
than  half  the  specimens  studied,  the  ventral  sur- 
face of  tail  with  low  bucklers  making  surface 
markedly  coarse  relative  to  smooth  shagreenlike 
texture  of  ventral  surface  of  disk.  All  fins  with 
at  least  a  few  tubercles;  dorsal  with  only  a  few 
tubercles  at  base,  but  pectoral,  ventral,  and  anal 
fins  with  tubercles  at  their  bases  and  running  out 
along  rays  for  a  third  to  three-fourths  their 
length.  Caudal  as  in  generic  description.  Skin 
covering  eyeballs  bearing  tiny  tubercles  with  rim 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


269 


around  cornea  beaded  by  a  row  of  small  buck- 
lers. Pectoral  membranes  thick,  opaque,  ventral 
surfaces  of  ends  of  rays  with  fleshy  pads. 

Color  in  preservative.  Ground  color  of  dorsal 
surface  of  body  light  tan  to  dark  brown  with 
round  dark  spots  (occasionally  rings)  in  an  elon- 
gate cluster  on  either  shoulder  and  in  a  band 
down  either  side  of  tail,  each  continuous  with  a 
cluster  in  axillae.  Sometimes  the  same  spotted 
pattern  occurs  on  face.  Ventral  surface  uniform- 
ly pale;  lips  pale.  The  small  rostrum  sometimes 
with  a  dusky  tip,  but  no  dark  subterminal  ring 
except  in  a  few  small  specimens;  iris  golden  or 
black.  Basal  portions  of  pectoral  fins  pale,  oc- 
casionally with  a  few  faint  spots,  darkening  dis- 
tally  so  that  frequently  a  distinct  black  border 
is  formed.  Ventrals  and  anal  the  same  pale 
ground  color  as  ventral  surface  of  body,  anal 
with  a  dusky  spot  distally  in  a  few  small  speci- 
mens. Dorsal  dusky  with  faint  splotches  or  sim- 
ply pale.  About  one-half  the  specimens  at  hand 
have  no  dark  markings  on  caudal,  but  remainder 
show  pattern  common  for  genus. 

Color  in  life.  The  following  description  from 
three  color  transparencies  of  a  freshly  killed 
specimen  captured  in  winter  off  North  Carolina; 
one  transparency  shows  a  dorsal,  one  a  lateral, 
and  one  a  ventral  view. 

Ground  color  of  dorsal  surface  of  body  me- 
dium brown  with  whitish  patches  where  skin 
evidently  abraded  from  protruding  tops  of  buck- 
lers. Darker  mottlings  of  pigment  in  axillae.  Fine 
black  lines  radiating  from  apices  of  large  buck- 
lers. An  elongate  cluster  of  round  dark  spots  on 
either  shoulder;  among  these  spots  are  sprinkled 
a  few  smaller,  intensely  orange  spots,  about  four 
on  each  side  (cast  shadows  on  the  photographs 
may  be  obscuring  other  orange  spots).  Pectoral 
and  caudal  fins  with  mottled  brownish  basal  por- 
tions fading  to  translucent  white  with  pale  tan 
striations  on  middle  portions  and  terminating  in 
broad,  bright  orangy-red  borders  blending  to 
pure  dark  red  and  finally  to  narrow  black  distal 
margins. 

Face  the  same  brownish  ground  color  as  dor- 
sal surface  of  body.  Lips  bright  orangy  red;  iris 
a  rosy-brass  color  with  tan  spots  in  a  circle 
around  pupil.  Ventral  operculum  well  extended 
in  example  and  the  same  rosy-brass  color  as  rest 
of  iris. 

Ventral  surface  of  body  with  a  transparent 
wash  of  pinkish  orange  over  a  pale  gray  surface. 


DISTRIBUTION  (Fig.  17). — Atlantic  coast  of 
the  United  States  from  Cape  Hatteras,  North 
Carolina,  to  the  Florida  Keys.  (One  specimen 
from  Jamaica,  USNM  160732,  150.0  mm  SL,  has 
the  general  shape  of  the  head  and  the  small 
mouth  that  characterize  rostellum,  but  it  differs 
from  rostellum  in  other  body  proportions  and 
has  no  diagnostic  color  markings;  I  am  unable 
to  assign  this  specimen.)  Bathymetric  range:  28- 
228m. 

NAME. — The  name  rostellum  is  the  diminu- 
tive of  the  word  rostrum,  referring  to  the  partic- 
ularly small  rostrum  in  this  species. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  For  data  from  COMBAT  stations  (  =  C)  and 
SILVER  BAY  stations  (  =  SB),  refer  to  Bullis  and  Thompson 
(1965);  for  data  from  PELICAN  stations(  =  P),  refer  to  Anderson 
(1956).  HOLOTYPE.  USNM  188809,  C-336,  Atlantic  coast  of 
Florida;  a  male  129.5  mm  SL. 

PARATYPES.  North  Carolina:  FMNH  64338  (1),  C-384; 
USNM  72288  (1),  Cape  Lookout;  USNM  1 1 1848  ( 1),  Beaufort; 
USNM  132093  (1),  ALBATROSS  sta.  2417,  33°18'30"N, 
77°07'W,  174  m:  USNM  148317  (2),  ALBATROSS  sta.  2600, 
34°39'40"N,  75°35'30"W.  159  m;  USNM  148320  (3),  ALBATROSS 
sta.  2602,  34°38'30"N,  75°33'30"W,  227  m;  USNM  151919  (1), 
ALBATROSS  III  sta.  35,  34°45.5-48'N,  75°28'-33'W,  77-88  m; 
USNM  151962  (1),  ALBATROSS  III,  31-B,  sta.  14,  33°26'N, 
77°03.5-06'W,  59  m;  USNM  151971  (1),  ALBATROSS  III,  31-D, 
sta.  3,  34°H'N,  76°06'W,  82-119  m.  South  Carolina:  CAS 
23925  (3),  SB- 1782;  FMNH  64100  (1),  BOWERS  sta.  50, 
3T50'N,  79°45'W,  46  m;  USNM  84510  (4),  ALBATROSS  sta. 
2312,  32°54'N,  77°53'30"W,  161  m.  Georgia:  ANSP  103634(1), 
P-178-10;  CAS-SU  62117  (2),  SB-1768;  FMNH  66385  (15), 
BOWERS  sta.  32,  30°14'N,  80°16'W,  73  m:  UF  24189  (1), 
Brunswick;  USNM  149959  (2);  USNM  188810  (1),  P-198-5. 
Florida:  FMNH  66388  (2),  SB-3442;  MCZ  32940  (1),  inlet  near 
New  Smyrna;  MCZ  45088  (3),  SB-3022;  MCZ  45089  (1),  SB- 
2775;  MCZ  45090(1),  P- 168-2:  MCZ  45091  (1),  Cape  Canaveral 
Pier;  MCZ  45092  (1),  SB-2771;  MCZ  45093  (1),  P- 169-3; 
USNM  18031  (1),  St.  Augustine;  USNM  38026  (1),  3.2  km  s 
of  Mosquito  Inlet,  Hillsborough  River:  USNM  131628  (2), 
ALBATROSS  sta.  2641,  25°11'30"N,  80°10'W,  110  m:  USNM 
188790  (1),  P-208-8:  USNM  188794  (1),  P-171-2:  USNM 
188806  (L),  P-169-8;  USNM  188807(1),  C-334:  USNM  188813 
(2),  C-336  (same  haul  as  holotype).  Florida  Straits:  CAS-SU 
62123  (1),  SB-2039:  UF  24190(1),  C-455.  Florida  Keys:  USNM 
117004  (2),  Tortugas;  USNM  117005  (1),  Tortugas. 

OTHER:  USNM  83785-83786  (3),  ALBATROSS  sta.  2311  and 
2318,  32°55'N,  77°54'W  (South  Carolina)  and  24°25'45"N, 
81°46'00"W  (Florida  Keys),  145  and  82  m. 


Ogcocephalus  declivirostris  new  species 

(Figure  24C-E) 

Ogcocephalus  sp.:  HILDEBRAND  1954:322  [off  Texas:  partial 
description:  stomach  contents];  WALLS  1975:120  [N  Gulf  of 
Mexico;  called  Ogcocephalus  sp.  B;  partial  description; 
figs.];  HOESE  AND  MOORE  1977:144,  fig.  100  [NW  Gulf  of 
Mexico;  key;  partial  description;  range]. 


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DIAGNOSIS  AND  COMPARISONS. — An  Ogco- 
cephalus  with  rostrum  generally  sloping  down- 
ward with  respect  to  the  long  axis  of  the  body 
instead  of  being  horizontal  or  tilted  upward.  O. 
declivirostris  differs  from  all  its  congeners  ex- 
cept pumilus  and  notatus  in  lacking  the  devel- 
opment of  fleshy  pads  on  the  ventral  surfaces  of 
the  pectoral  and  pelvic  rays.  It  may  be  separated 
from  the  long-nosed  species,  pumilus,  corniger, 
and  vespertilio,  by  the  length  of  the  rostrum 
(length  of  rostrum  into  length  of  disk  margin  3.4- 
8.4  in  declivirostris,  1.5-3.0  in  the  long-nosed 
species,  Fig.  9)  and  from  notatus  by  the  width 
of  the  mouth  (width  of  mouth  into  length  of  disk 
margin  2.8-3.7  in  declivirostris,  1.8-2.5  in  no- 
tatus). It  is  distinguished  from  all  species  except 
pumilus,  parvus,  and  corniger  by  its  low  pec- 
toral ray  count  (usually  11  in  declivirostris,  12 
or  more  in  other  species  except  pumilus,  par- 
vus, and  corniger,  Table  3).  O.  declivirostris  is 
sympatric  with  O.  pantostictus,  both  being 
species  of  the  northern  and  western  coasts  of 
the  Gulf  of  Mexico;  besides  the  lower  pectoral 
fin  ray  count  in  declivirostris,  just  mentioned, 
declivirostris  has  a  lower  count  for  the  sub- 
opercular  lateral-line  series  than  pantostictus 
(modally  7  in  declivirostris,  9  in  pantostictus). 
The  range  of  parvus  overlaps  with  that  of  de- 
clivirostris in  the  northern  Gulf  of  Mexico;  de- 
clivirostris is  distinguished  from  parvus  by  its 
flatter  head  (head  depth  into  SL  4.3-5.1  in  de- 
clivirostris, 3.1-4.2  in  parvus)  and  larger  mouth 
(width  of  mouth  into  head  depth  1.4-1.9  in  de- 
clivirostris, 2.0-2.8  in  parvus). 

DESCRIPTION. — Counts  and  measurements 
from  99  specimens,  39.6  to  137.0  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6;  in  follow- 
ing description,  counts  for  holotype  given  in 
brackets.  In  declivirostris  all  counts  except  ver- 
tebral count  at  low  end  of  range  of  variation 
within  genus.  Pectoral  fin  ray  count  usually  11 
[11,11];  range  9-12.  Subopercular  lateral-line 
scale  count  usually  8  [8,9];  range  7-9.  Mean  for 
lateral-line  scale  count  25  [25,27];  range  22-28. 
Vertebral  count  most  frequently  20;  range  18- 
20.  [D.  5;  A.  3.] 

Proportions.  Proportions  expressed  as  ratios 
given  by  the  mean  followed  by  the  range  in  pa- 
rentheses and  the  proportion  for  the  holotype  in 
brackets. 

Length  of  disk  margin  intermediate  relative  to 
range  of  variation  for  genus,  2.4(2.3-2.6)[2.4]  in 


standard  length.  Tail  stout  and  broad,  its  width 
1.6(1.3-2.0)[1.3]  in  length  of  disk  margin,  taper- 
ing slightly  but  rounding  off  more  abruptly  at 
caudal  fin  than  in  other  species.  Caudal  peduncle 
thick,  its  depth  2.5(2.3-2.8)[2.5]  in  head  depth. 
Rostrum  a  rod-shaped  protuberance  with  a  nar- 
row base,  generally  pointing  downward  but 
sometimes  horizontal  relative  to  long  axis  of 
body;  never  tilted  upward.  Length  of  rostrum 
5.7(3.4-8.4)[7.0]  in  length  of  disk  margin;  the 
series  at  hand  shows  that  during  ontogeny  the 
rostrum  becomes  relatively  shorter.  Aperture  of 
illicial  cavity  approximately  oval  to  subtriangu- 
lar,  wider  than  high  in  medium-  and  large-sized 
specimens,  higher  than  wide  in  specimens  less 
than  about  65  mm  SL.  Cranium  when  viewed 
from  front  rising  in  a  gradual  curve  above  disk, 
head  depth  1.9(1.8-2.1)[1.9]  in  length  of  disk 
margin.  Width  of  cranium  3.2(2.7-3.7)[3.3]  in 
length  of  disk  margin.  The  following  in  width  of 
cranium:  eye  1.4(1.1-1.6)[1.6],  lateral  ethmoid 
width  1.6(1. 3- 1.9)  [1.6].  Interorbital  space  con- 
cave, narrow,  its  width  2.7(2.1-3.4)[3.2]  in  width 
of  cranium,  4.5(3.3-5.7)[5.7]  in  head  depth. 
Mouth  average  size,  its  width  1.6(1.4-1.9)[1.5] 
in  head  depth,  3.1(2.8-3.7)[2.8]  in  length  of  disk 
margin;  length  of  jaw  2.4(2. 1-2. 7)[2.1]  in  head 
depth.  Upper  lip  moderately  fleshy,  of  an  even 
width  for  its  entire  length;  lower  lip  thickened 
medially,  the  thickening  becoming  a  pronounced 
lobe  in  largest  specimens.  Dorsal  fin  relatively 
large,  its  length  2.8(2.4-3. 4)[2. 8]  in  length  of  disk 
margin;  anal  fin  also  large,  its  length  2.5(2.1- 
2.9)[2.6]  in  length  of  disk  margin. 

Integument.  Dermal  cirri  well  developed  on 
lateral  sides  of  tail  and  disk,  more  so  than  in 
other  species  of  Ogcocephalus.  Cirri  present  on 
tip  of  rostrum  in  small  specimens.  Large  buck- 
lers arranged  as  described  in  generic  descrip- 
tion. Dorsal  fin  devoid  of  tubercles,  anal  fin  with 
only  a  few  at  base.  Pectoral  fins  with  tubercles 
at  base  on  both  dorsal  and  ventral  surfaces,  a 
few  tubercles  extending  out  along  anterior  and 
posterior  rays.  Pelvics  with  tubercles  present  on 
basal  quarter  of  ventral  surface  but  none  on  dor- 
sal surface.  Caudal  as  in  generic  description. 
Skin  covering  eyeballs  bearing  tiny  tubercles 
with  rim  around  cornea  beaded  by  a  row  of  small 
bucklers  in  small  specimens,  disappearing  in 
large  specimens.  Pectoral  membranes  relatively 
thin  and  translucent;  no  fleshy  pads  on  ventral 
tips  of  rays. 

Color  in  preservative.  Ground  color  of  dorsal 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


271 


•    parvus 
©   pumilus 
o    declivirostris 


FIGURE  25.     Localities  for  study  material  of  Ogcocephalus  parvus,  O.  pumilus,  and  O.  declivirostris.  O.  parvus  also  known 
from  Recife,  Brazil  (Gunther  1880). 


surface  of  body  gray  or  tan,  apices  of  bucklers 
paler  than  ground  color.  Dim  rounded  spots 
sometimes  clustered  on  shoulders,  face,  in  ax- 
illae, and  along  sides  of  tail,  occurring  mostly  in 
small  specimens.  A  few  small  specimens  have 
these  spots  covering  entire  disk  dorsally.  Ven- 
tral surface  of  body  uniformly  pale;  lips  pale, 
occasionally  with  dusky  splotches.  Rostrum 
with  a  diffuse  black  tip  in  small  specimens,  this 
mark  disappearing  in  larger  specimens.  Iris 
black  or  golden,  sometimes  golden  with  a  ring 
of  dim  spots  circling  pupil.  Pectoral  fins  dusky, 
sometimes  darkened  towards  tips,  but  no  pro- 
nounced dark  borders  except  in  some  small  in- 
dividuals. Pelvics  and  anal  the  same  pale  ground 
color  as  ventral  surface  of  body,  anal  sometimes 
with  a  dusky  tip  in  small  specimens.  Dorsal 
dusky  with  inconspicuous  splotches.  Caudal  as 
described  in  generic  description  except  for  a  few 
specimens  that  have  the  caudal  uniformly 
dusky. 

Color  in  life.  Unknown. 

DISTRIBUTION  (Fig.  25). — Northern  and  west- 
ern coasts  of  Gulf  of  Mexico  from  longitude 
86°W  to  latitude  26°N.  One  record  from  Florida 
Straits.  Bathymetric  range:  3.5-180  m  except  for 
Florida  Straits  record  (388  m). 

NAME. — The  name  Ogcocephalus  decliviros- 
tris means  an  Ogcocephalus  with  rostrum  slop- 
ing downward. 


MATERIAL  EXAMINED. — HOLOTYPE:  FMNH  64136,  off 
Texas,  28°02'N,  96°03'W,  40  m;  a  male  129.3  mm  SL. 

PARATYPES.  Numbers  in  parentheses  are  numbers  of  spec- 
imens. For  data  from  OREGON  stations  (  =  O),  refer  to  Springer 
and  Bullis  (1956)  and  Bullis  and  Thompson  (1965).  Gulf  coast 
of  Florida:  ANSP  103628  (1),  O-331;  FMNH  61579  (1),  O-858; 
FMNH  64120  (2),  O-944.  Florida  Straits:  UMML  3897  (1), 
O-1550.  Louisiana:  CAS  23930  (1),  O-2700;  CAS  23931  (1), 
O-2704;  CAS-SU  62119  (3),  PELICAN  sta.  84-3,  28°14'N, 
91°41'W,  55  m;  FMNH  46735  (1),  0-72;  FMNH  46736  (3), 
O-75;  FMNH  46737  (3),  O-283-284;  GCRL  557  (2)  and  GCRL 
559  (1),  s  of  Grand  Isle,  37  m;  GCRL  558  (2)  and  GCRL  560 
(1),  s  of  Grand  Isle,  27  m;  MCZ  45072  (3),  PELICAN  sta.  74-3, 
29°04'N,  88°44.5'W,  110m;  MCZ  45073(1),  PELICAN  sta.  72-6, 
29°11.5'N,  88°52'W,  73  m;  MCZ  45074(1),  PELICAN  sta.  69-6, 
28°48'N,  89°51'W,  55  m;  USNM  131166  (21),  Morgan  City, 
3.7-9.1  m;  USNM  188761  (1),  sw  of  Eugene  I.  Beacon, 
Atchafalaya  Bay;  USNM  188764  (1),  PELICAN  sta.  85-4, 
28°11'N,  91°24.5'W,  86  m;  USNM  188765  (1),  PELICAN  sta. 
85-3,  28°09'N,  91°27'W,  91  m;  USNM  188767  (1),  PELICAN 
sta.  81-8,  28°50'N,  91°29.5'W,  18  m;  USNM  188771  (1)  and 
USNM  188774  (1),  ssw  of  Eugene  I.  Beacon,  Atchafalaya 
Bay;  USNM  188777  (1),  PELICAN  sta.  86-3,  28°28'N, 
9P13.5'W,  40  m;  USNM  188784  (2),  PELICAN  sta.  34,  28°33'N, 
90°55'W,  26  m;  USNM  188792  (2),  PELICAN  sta.  84-1, 
28°22.5'N,  91°44.5'W,  59  m;  USNM  188796  (1),  PELICAN  sta. 
94-1,  28°27'N,  92°14'W,  53  m;  USNM  188799  (1),  PELICAN 
sta.  93-8,  28°33'N,  92°15.5'W,  44  m;  USNM  188801  (2),  PEL- 
ICAN sta.  93-3,  28°55'N,  92°15.5'W,  26  m.  Texas:  CAS-SU 
62140  (30),  Galveston;  FMNH  61576  (1),  27°04'N,  96°23'W, 
55  m;  FMNH  61577  (2),  28°02'N,  96°04'W,  40  m;  FMNH 
61578  (1),  28°02'N,  96°04'W,  37  m;  FMNH  61580  (1),  27°22'N, 
96°34'W,  49  m:  FMNH  61582  (1),  27°30'N,  96°28'W,  73  m; 
FMNH  61583  (1),  27°39'N,  96°28'W,  68  m;  FMNH  64135  (1), 
27°49'N,  96°31'W,  40  m;  FMNH  64137  (1),  27°34'N,  96°43'W, 
42  m;  FMNH  64138  (2),  off  Mustang  I.,  46-48  m;  USNM 


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FIGURE  26.  (A)  Paratype  of  Ogcocephalus  corniger  new  species,  61.0  mm  SL,  UF  24183,  from  w  of  Andros  I.,  Bahamas, 
24°25'N,  79°13'W,  showing  color  pattern  of  juvenile.  (B)  and  (C)  Ogcocephalus  corniger  new  species,  holotype,  124.5  mm  SL, 
USNM  188808,  Gulf  of  Mexico,  29°00'-10'N,  85°01'W.  (D)  and  (E)  Ogcocephalus  pumilus  new  species,  holotype,  57.5  mm 
SL,  USNM  188812,  Surinam,  7°05'N,  54°08'W. 


118606  (23),  Galveston;  USNM  171763  (2),  near  Brownsville; 
USNM  188776  (1),  SILVER  BAY  sta.  200,  27°59'N,  95°15'W, 
79-84  m.  Material  without  exact  locality:  USNM  188766  (1), 
Madame  Canaia. 

Ogcocephalus  pumilus  new  species 

(Figure  26D-E) 

DIAGNOSIS  AND  COMPARISONS. — A  long-nosed 
Ogcocephalus  distinctive  for  its  small  size  and 
lack  of  pigment  markings.  Ogcocephalus  pumi- 
lus is  the  smallest  member  of  the  genus;  maxi- 
mum standard  lengths  for  specimens  seen  in  this 
study  are  all  under  70  mm  (Table  1).  O.  pumilus 
differs  from  all  its  congeners  except  decliviros- 
tris  and  notatus  in  lacking  the  fleshy  ventral 
pads  on  the  tips  of  the  rays  of  the  pectoral  and 
pelvic  fins. 

Ogcocephalus  pumilus  is  sympatric  with  par- 


vus  and  nasutus,  but  as  it  does  not  occur  on  the 
coasts  of  the  southeastern  United  States  and 
Gulf  of  Mexico,  its  range  is  less  extensive  than 
that  of  parvus.  It  is  distinguished  from  parvus 
by  its  long  rostrum  (length  of  rostrum  into  length 
of  disk  margin  2.0-3.0  in  pumilus,  3.2-5.0  in 
parvus)  and  shorter  disk  margin  (length  of  disk 
margin  into  SL  2.2-2.7  in  pumilus,  1.9-2.2  in 
parvus)  as  well  as  its  pale  color  contrasted  with 
the  mottlings  and  spotted  patterns  seen  in  par- 
vus. O.  pumilus  is  also  distinguished  from  most 
examples  of  nasutus  by  its  lack  of  color  pattern 
as  well  as  its  diminutive  size  (nasutus  being 
among  the  largest  species  in  the  genus).  Also, 
many  specimens  of  pumilus  may  be  separated 
from  nasutus  by  the  pectoral  fin  ray  count  (mod- 
ally  1 1  in  pumilus,  12  or  13  in  nasutus,  Table  3). 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


273 


Off  the  northern  coast  of  South  America,  the 
range  of  pumilus  overlaps  that  of  notatus.  The 
two  species  are  separable  on  the  basis  of  the 
pectoral  fin  ray  count  (11-12  in  pumilus,  13-14, 
rarely  12,  in  notatus)  and  subopercular  lateral- 
line  scale  count  (range  5-7,  modally  6,  in  pumi- 
lus, 7-10,  modally  9,  in  notatus).  O.  pumilus 
also  has  a  markedly  smaller  mouth  than  notatus 
(width  of  mouth  into  length  of  disk  margin  2.6- 
3.4  in  pumilus,  1.8-2.5  in  notatus). 

DESCRIPTION. — Counts  and  measurements 
from  63  specimens,  25.8  to  66.7  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6.  In  fol- 
lowing description,  counts  for  holotype  given  in 
brackets.  All  counts  in  pumilus  low  compared 
to  counts  for  genus  as  a  whole.  Pectoral  fin  ray 
count  most  often  11  [11,11];  range  10-12.  Sub- 
opercular lateral-line  scale  count  most  often  6 
[6,7];  range  4-7.  Cheek  lateral-line  scale  count 
most  often  8  [8,8];  range  8-9.  Mean  for  lateral- 
line  scale  count  21  [21,21];  range  19-24.  Anal  fin 
ray  count  3  more  than  half  the  time  [3],  remark- 
able because  count  normally  4  throughout  fam- 
ily. [D.  4.]  Vertebral  counts  on  14  specimens  all 
19. 

Proportions.  Proportions  expressed  as  ratios 
given  as  mean  followed  by  range  in  parentheses 
and  proportion  for  holotype  in  brackets. 

Length  of  disk  margin  2.3(2.2-2.5)[2.3]  in  SL. 
Tail  moderately  wide,  its  width  1.8(1.6-2. 1)[1. 8] 
in  length  of  disk  margin.  Caudal  peduncle 
2.8(2.5-3.2)[2.6]  in  head  depth.  The  grotesque 
rostrum  thin  and  long,  its  length  2.4(2.0- 
3.0)[2.4]  in  length  of  disk  margin,  its  long  axis 
about  horizontal  with  body  axis  rather  than  tilt- 
ed upward.  Aperture  of  illicial  cavity  subtrian- 
gular,  higher  than  wide.  Head  depth  1.9(1.7- 
2.1)[2.1]  in  length  of  disk  margin.  Width  of  cra- 
nium 3.1(2.7-3.5)[3.3]  in  length  of  disk  margin. 
The  following  in  width  of  cranium:  eye  1.3(1.1- 
1.5)[1. 3];  lateral  ethmoid  width  1.7(1.5-1. 9)[1.6]. 
Interorbital  space  slightly  concave,  narrow,  its 
width  3.1(2.4-3.7)[2.8]  in  width  of  cranium, 
5.1(3.8-6.2)[4.4]  in  head  depth.  Mouth  moder- 
ate-sized, its  width  1. 5(1.4-1. 7)[1. 7]  in  head 
depth,  3.0(2.7-3.4)[3.2]  in  length  of  disk  margin; 
length  of  jaw  2. 4(2. 1-2. 6) [2. 4]  in  head  depth. 
Upper  lip  moderately  fleshy,  of  an  even  width 
for  its  entire  length;  lower  lip  fleshy,  no  median 
lobe.  Dorsal  and  anal  fins  about  average  in  pro- 
portion to  body  size,  length  of  dorsal  fin  3.3(2.3- 


4.3)[3.9]  in  length  of  disk  margin;  length  of  anal 
fin  2.9(2.4-3. 7)[3.0]  in  length  of  disk  margin. 

Integument.  Dermal  cirri  usually  present  on 
lateral  sides  of  tail  and  disk  margin.  No  cirri  on 
tip  of  rostrum.  Large  bucklers  arranged  as  de- 
scribed for  genus.  Dorsal  fin  devoid  of  tubercles, 
anal  fin  with  only  a  few  near  base.  Pectorals  and 
pelvics  with  tubercles  at  base  and  running  out 
on  the  fins  for  1A  to  Vi  their  length.  Caudal  as  in 
generic  description.  Skin  covering  eyeballs 
bearing  tiny  tubercles  with  rim  around  cornea 
often  beaded  by  a  row  of  small  bucklers.  Pec- 
toral membranes  thin,  translucent;  pectoral  rays 
without  fleshy  pads  on  their  ventral  tips. 

Color  in  preservative.  Dorsal  surface  of  body 
white  to  pale  gray  or  grayish  brown;  in  the  latter 
case,  apices  of  bucklers  paler  than  ground  color. 
Pigment  markings  usually  absent;  if  present, 
they  are  dusky  and  ill-defined  spots,  occurring 
on  shoulders,  lateral  sides  of  tail,  and  (very  in- 
frequently) face.  Ventral  surface  of  body  uni- 
formly pale;  lips  also  pale.  Rostrum  with  a  dusky 
subterminal  ring.  Iris  black  or  silvery.  Pectorals 
colorless  in  white  examples,  dusky  in  gray  ones; 
in  the  latter,  the  gray  color  may  shade  to  a  dark- 
er tone  towards  tips  of  pectorals.  Pelvics  and 
anal  the  same  pale  whitish  color  as  ventral  sur- 
face of  body;  anal  sometimes  with  a  dusky  tip. 
Dorsal  colorless  in  white  specimens,  dusky  in 
others,  sometimes  with  a  faintly  darker  stripe  or 
two.  Caudal  colorless  to  dusky  in  white  exam- 
ples; gray  or  brownish  individuals  with  caudal 
as  in  generic  description. 

Color  in  life.  From  a  color  transparency  taken 
shortly  after  capture:  body  pale  bluish  gray, 
ventral  surface  of  body  paler  than  dorsal  sur- 
face, the  whole  suffused  with  a  wash  of  trans- 
lucent pinkish  orange.  Rostrum  a  deeper  orange. 
Distal  thirds  of  pectorals  and  caudal  deep  or- 
ange. 

DISTRIBUTION  (Fig.  25). — Known  from  the 
Bahamas  just  north  of  Cuba,  from  Puerto  Rico, 
the  Leeward  Islands,  the  western  Caribbean, 
and  the  northern  coast  of  South  America  to  Sur- 
inam. Bathymetric  range:  35-348  m. 

NAME. — The  name  pumilus,  which  is  derived 
from  a  word  meaning  dwarf,  is  given  in  refer- 
ence to  the  diminutive  size  and  grotesque  ap- 
pearance of  this  species. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  For  data  from  OREGON  (  =  O)  and  SILVER 


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BAY  (=SB)  stations,  refer  to  Bullis  and  Thompson  (1965). 
HOLOTYPE:  USNM  188812,  Surinam,  O-2018,  57.5  mm  SL. 
PARATYPES.  Northern  Bahamas:  USNM  188759  (2),  SB- 
2455.  Puerto  Rico:  FMNH  64843  (2)  and  FMNH  64844  (10), 
O-2626;  FMNH  64845  (2),  O-2627;  MCZ  45085  (3),  O-2668; 
MCZ  45087  (8),  O-2626.  Virgin  Islands:  CAS  23920  (1),  O- 
2618;  FMNH  64841  (4),  O-2616;  USNM  108353  (1), 
18°25'30"N,  63°32'36"W.  Leeward  Islands:  FMNH  64846  (5), 
O-2631;  FMNH  64848  (2),  O-2633;  FMNH  64891  (33),  O-2231; 
UMML  7488  (14),  O-2632;  USNM  190467  (1),  O-2632.  Hon- 
duras: CAS  23921  (1),  O-1865.  Guyana:  ANSP  103632  (3),  O- 
2231;  ANSP  103633  (1),  O-2262;  CAS  23919  (1),  O-2249;  CAS- 
SU  62122  (3),  O-2345;  CAS-SU  62125  (1),  O-2259;  FMNH 
64893  (3),  O-2249;  FMNH  65965  (1),  O-2001;  FMNH  65966 
(1),  O-2234;  FMNH  65967  (3),  O-2248;  FMNH  65968  (1),  O- 
2250;  FMNH  65969  (3),  O-2261;  FMNH  65970  (4),  O-2344; 
MCZ  45086  (3),  O-2261;  USNM  185984  (3),  8°33'N,  58°46'W. 
Surinam:  FMNH  64899  (2),  O-2330. 

Ogcocephalus  corniger  new  species 

(Figure  264 -C) 

Ogcocephalus  vespertilio  (non  Linnaeus):  LONGLEY  AND  HIL- 
DEBRAND  1941:311,  314  [Tortugas,  Florida;  key;  description]. 

Ogcocephalus  sp.:  LONGLEY  AND  HILDEBRAND  1941:311,  315 
[Tortugas,  Florida;  key;  partial  description  by  Hildebrand]; 
WALLS  1975: 1 18  [northern  Gulf  of  Mexico;  partial  descrip- 
tion; figs.;  called  Ogcocephalus  sp.  A;  color  photograph]. 

Oncocephalus  vespertilio  (non  Linnaeus):  FOWLER  1952:14 
[off  southern  Florida;  partial  description]. 

Probable  references: 

Lophius  nostratus  MITCHILL  1818:325  [''Straits  of  Bahamas"; 
a  misspelling  of  Lophius  rostratus  Shaw]. 

Malthe  vespertilio  (non  Linnaeus):  UHLER  AND  LUGGER 
1876:77  [southern  Chesapeake  Bay;  partial  description]. 

Oncocephalus  vespertilio  (non  Linnaeus):  GARMAN  1896:87 
[off  Key  West,  Florida;  partial  description]. 

Ogcocephalus  vespertilio  (non  Linnaeus):  H.  M.  SMITH 
1907:401,  figs.  185-186  [off  Beaufort,  North  Carolina;  par- 
tial description;  figures  the  same  as  Jordan  and  Evermann 
1898:pl.  392,  figs.  958  and  958b  (legend:  Ogcocephalus  ves- 
pertilio)]. 

DIAGNOSIS  AND  COMPARISONS. — A  long-nosed 
Ogcocephalus  distinctive  for  its  dorsal  color 
pattern,  which  consists  of  small  pale  round  spots 
close  together  and  evenly  distributed  over  the 
darker  background  color  of  the  dorsal  surface  of 
the  body  (specimens  under  about  65  mm  SL  lack 
this  pattern;  see  sections  on  color  below). 

The  long  rostrum  in  corniger  separates  it  from 
all  other  species  of  Ogcocephalus  except  ves- 
pertilio, pumilus,  long-nosed  morphs  of  nasutus, 
and  small  examples  of  cubifrons  less  than  50 
mm  SL  (Fig.  9).  The  range  of  corniger  (coasts 
of  southeastern  United  States  and  eastern  Gulf 
of  Mexico)  is  widely  separated  from  that  of  ves- 
pertilio (coast  of  Brazil)  and,  at  this  time,  ap- 
pears to  be  separated  from  that  of  pumilus 
(northern  Bahamas,  Caribbean,  and  coast  of 
South  America  to  Surinam).  Besides  separate 


ranges,  other  aspects  separate  these  long-nosed 
species.  O.  corniger  differs  from  vespertilio  in 
its  lower  number  of  pectoral  rays  (10-12  in  cor- 
niger, 13-15  in  vespertilio)  and  lower  number  of 
subopercular  lateral-line  scales  (usually  5-7  in 
corniger,  usually  8-9  in  vespertilio,  Table  6);  it 
differs  from  pumilus  in  its  larger  body  size 
(pumilus  is  less  than  70  mm  in  SL)  and  in  having 
fleshy  pads  on  the  ventral  tips  of  the  pectoral 
rays  which  pumilus  lacks. 

The  range  of  corniger  overlaps  the  ranges  of 
nasutus  and  cubifrons,  but  long-nosed  forms  of 
nasutus  and  cubifrons  may  be  distinguished 
from  corniger  by  the  pectoral  fin  ray  count 
(modally  11  in  corniger,  usually  12  or  13  in  na- 
sutus and  cubifrons,  Table  3).  O.  corniger  dif- 
fers further  from  nasutus  in  the  number  of  cheek 
lateral-line  scales  (usually  8  in  corniger,  modally 
9  in  nasutus,  Table  5)  and  from  cubifrons  by  its 
longer  jaw  (length  of  jaw  into  width  of  cranium 
1.2-1.5  in  corniger,  1.6-2.0  in  cubifrons). 

Two  short-nosed  species  of  Ogcocephalus, 
parvus  and  rostellum,  occur  within  the  range  of 
corniger  but  are  separable  from  corniger  not 
only  by  the  proportions  of  the  rostrum  but  also 
by  color  pattern — neither  ever  has  the  pattern 
of  small  round  pale  spots  on  a  darker  back- 
ground seen  in  corniger.  Other  characters  that 
separate  corniger  from  parvus  include  the 
smoother  shagreenlike  quality  of  the  integument 
in  corniger  compared  to  the  rough,  lumpy  sur- 
face of  the  integument  in  parvus,  and  the  larger 
mouth  in  corniger  (width  of  mouth  into  head 
depth  1.5-2.0  in  corniger,  2.0-2.8  in  parvus). 
An  additional  character  separating  corniger 
from  rostellum  is  the  pectoral  fin  ray  count 
(modally  1 1  in  corniger,  modally  13  in  rostellum, 
Table  3). 

DESCRIPTION. — Counts  and  measurements 
from  57  specimens  26.2  to  134.0  mm  SL  (Table 

1). 

Counts.  Counts  given  in  Tables  2-6.  In  the 
following  description,  counts  for  holotype  given 
in  brackets.  Counts  in  corniger  low  compared 
to  those  of  genus  as  a  whole.  Pectoral  fin  ray 
count  usually  11  [11,11];  range  10-12.  Suboper- 
cular lateral-line  scale  count  usually  6  or  7  [6,6]; 
range  4-8.  Cheek  lateral-line  scale  count  8  [8,8] 
(the  commonest  count  for  genus);  range  7-9. 
Mean  for  lateral-line  scale  count  nearly  21 
[20,20];  range  18-24.  Majority  of  sample  of  cor- 
niger had  19  vertebrae,  the  most  usual  vertebral 
count  for  the  genus.  [D.  3;  A.  4.] 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


275 


Proportions.  Proportions  expressed  as  ratios 
given  as  mean  followed  by  range  in  parentheses 
and  proportion  for  holotype  in  brackets. 

Length  of  disk  margin  intermediate  relative  to 
range  of  variation  in  genus,  2.3(2.1-2.5)[2.4]  in 
SL.  Tail  narrow  to  moderately  wide,  the  width 
2.0(1.7-2.4)[1.7]  in  length  of  disk  margin,  taper- 
ing to  a  relatively  thin  caudal  peduncle;  depth 
of  caudal  peduncle  3.3(2.8-4.1)[3.3]  in  head 
depth.  Rostrum  with  a  wide  base  but  becoming 
slender  distally,  upturned  and  very  long,  the 
length  2.0(1. 7-2.3)[2.1]  in  length  of  disk  margin. 
Aperture  of  illicial  cavity  triangular,  always 
higher  than  wide.  Cranium,  when  viewed  from 
front,  rising  steeply  above  disk;  head  relatively 
deep,  its  depth  1.8(1.6-1.9)[1.7]  in  disk  margin. 
Width  of  cranium  3.3(2.9-3.5)[3.2]  in  length  of 
disk  margin.  Eye  comparably  large,  1.3(1.1- 
1.7) [1.5]  in  width  of  cranium.  Lateral  ethmoid 
width  1.7(1.5-2.0)[1.7]  in  width  of  cranium.  In- 
terorbital  space  flat  to  moderately  convex,  its 
width  2.1(1.7-2.5)[1.9]  in  width  of  cranium, 
3.9(3. 1^.6)[3.6]  in  head  depth  (except  one  spec- 
imen, which  has  these  proportions  2.9  and  5.7, 
respectively).  Mouth  average  size,  its  width 
1.6(1.5-2.0)[1.5]  in  head  depth,  2.9(2.6-3. 7)[2.6] 
in  length  of  disk  margin;  length  of  jaw  2.4(2.2- 
2.6)[2.5]  in  head  depth.  Upper  lip  fleshy,  of  an 
even  width  for  its  entire  length;  lower  lip  also 
fleshy,  thickly  swollen  medially.  Dorsal  fin  small 
compared  to  other  species  in  genus,  its  length 
4.8(3.7-5.8)[5.8]  in  length  of  disk  margin;  anal 
fin  also  small,  its  length  3.0(2.5-3.5)[3.0]  in 
length  of  disk  margin. 

Integument.  Dermal  cirri  absent,  or  at  least 
inconspicuous,  except  near  tip  of  rostrum, 
where  fairly  long  cirri  occur  in  subterminal  ring. 
Large  bucklers  arranged  as  in  generic  descrip- 
tion. Dorsal  fin  devoid  of  tubercles;  anal  fin  with 
tiny  tubercles  covering  proximal  third  or  half  of 
fin.  Pectorals  and  pelvics  with  tiny  tubercles  at 
base  on  both  dorsal  and  ventral  surfaces,  run- 
ning out  along  fin  rays  for  a  third  to  half  their 
length.  Caudal  as  in  generic  description.  Skin 
covering  eyeballs  bearing  fine  tubercles  with  rim 
around  cornea  beaded  by  a  row  of  slightly  larger 
ones.  Pectoral  membranes  thick,  opaque,  ven- 
tral surfaces  of  the  ray  tips  with  well-developed 
fleshy  pads  (pads  not  developed  in  small  speci- 
mens). 

Color  in  preservative.  Dorsal  surface  of  body 
uniformly  covered  with  small  pale  spots  on  a 
darker  ground  color,  the  ground  color  generally 


FIGURE  27.     Localities  for  study  material  of  Ogcocephalus 
corniger. 


brown  or  gray,  becoming  darker  on  shoulders  in 
many  examples;  sometimes  spots  so  close  to- 
gether that  ground  color  may  appear  as  a  retic- 
ulum.  The  pale  spots  are  unpigmented  apices  of 
bucklers.  Specimens  under  about  65  mm  SL 
generally  lack  this  color  pattern  and  are  uni- 
formly pale  or  dusky,  but  occasionally,  dark 
rings  occur  clustered  on  shoulders.  Ventral  sur- 
face of  body  uniformly  pale;  lips  pale.  Rostrum 
with  a  diffuse  dark  subterminal  ring,  incomplete 
below.  Iris  black  or  golden,  or  golden  with 
spots,  but  never  with  radiating  lines.  Pectorals 
pale  to  dusky  proximally  with  broad  dark  brown 
or  black  margins;  ray  tips  white,  ventral  pads  of 
rays  white.  Pelvic  and  anal  fins  the  same  ground 
color  as  ventral  surface  of  body;  pelvics  some- 
times with  darkened  or  blotched  tips  in  dark 
specimens;  anal  most  often  with  distal  third  very 
dark  to  black.  Dorsal  dusky,  often  with  a  pale 
diagonal  stripe  or  two.  Caudal  as  in  generic  de- 
scription, the  border  always  dark  to  black. 

Color  in  life.  The  following  description  from 
three  color  transparencies  made  of  a  freshly 
killed  specimen  captured  in  winter  off  North 
Carolina.  One  transparency  shows  a  dorsal  view 
of  the  fish  and  one  a  ventral  view;  the  remaining 
one  shows  a  lateral  view  of  the  head. 


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Ground  color  of  dorsal  surface  of  body  choc- 
olate brown  with  fine,  close-set  pearl-gray  spots 
completely  covering  the  surface,  including  face 
and  skin  covering  eyeballs.  Even  iris  of  eye  has 
the  spotted  pattern,  with  a  brassy  highlight 
around  pupil.  Lips  bright  red-orange,  with  a 
translucent  wash  of  red-orange  pigment  over 
lower  face  and  lower  edge  of  disk,  where  it  be- 
comes continuous  with  the  same  red-orange 
wash  over  ventral  surface  of  body,  including 
pelvic  and  anal  fins.  This  red-orange  coloration 
on  ventral  surface  is  more  intense  on  chin,  on 
a  ring  encircling  anus,  and  on  tail  including  anal 
fin,  which  also  has  a  black  tip.  Pectoral  fins  with 
translucent  red-orange  color  basally,  the  rays 
becoming  deep  red-orange  distally  and  the  fins 
each  with  a  broad  blackish  margin.  Caudal  dark 
basally,  becoming  pale  red-orange  in  middle 
portion  with  rays  a  darker  shade  than  mem- 
branes, and  black  again  distally  in  a  broad  mar- 
gin. 

Longley  (Longley  and  Hildebrand  1941)  stat- 
ed for  examples  from  off  Tortugas  (his  Ogco- 
cephalus  vespertilio)  that  the  pectorals  and  cau- 
dal were,  "buff-yellow  over  basal  half,  passing 
through  maroon-purple  to  prune  purple  at  the 
tips,  the  contrast  between  the  darker  outer  third 
and  the  yellow  basal  two-thirds  being  sharper  on 
the  under  side  of  the  pectorals;  belly,  maxillary 
membranes  and  buccal  cavity  coppery  red." 
Walls  (1975)  presents  a  photograph  of  an  ex- 
ample from  the  Gulf  of  Mexico  (his  Ogcoceph- 
alus  sp.  A)  which  shows  the  pectorals  red-or- 
ange basally,  succeeded  by  bright  yellow,  which 
gives  way  to  more  red-orange,  which  in  turn 
gives  way  to  black  borders. 

DISTRIBUTION  (Fig.  27). — Known  from  the 
Atlantic  coast  of  the  United  States  from  Cape 
Lookout,  North  Carolina,  to  Florida,  Campeche 
Banks,  and  the  northeastern  Gulf  of  Mexico  to 
Louisiana;  one  record  from  off  Great  Inagua  I., 
Bahamas.  Bathy metric  range:  29-230  m. 

NAME. — This  form  is  named  corniger,  mean- 
ing horn-bearer,  in  reference  to  the  long,  up- 
turned rostrum. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  For  data  from  early  OREGON  stations  (  = 
O),  refer  to  Springer  and  Bullis  (1956):  for  data  from  later 
OREGON  stations  and  for  COMBAT  (=C)  and  SILVER  BAY  (= 
SB)  stations,  refer  to  Bullis  and  Thompson  ( 1965).  HOLOTYPE: 
USNM  188808,  Gulf  coast  of  Florida,  O-729-730;  a  male  124.5 
mm  SL. 

PARATYPES.   North   Carolina:   CAS   23927  (1),   SB-2930: 


USNM  152030  (1).  Atlantic  coast  of  Florida:  ANSP  103626  (1), 
C-I01;  CAS-SU  62124  (1),  C-497;  CAS-SU  62128(1),  SB-1931; 
USNM  161375  (1),  Palm  Beach.  Bahamas:  FMNH  66387  (1), 
SB-3496;  UF  24183  (1),  SB-2470.  Florida  Straits:  CAS  23926 
(1),  SB-437;  UF  24182  (1),  SB-2362.  Florida  Keys:  FMNH 
64124  (6),  O-1020;  USNM  37853  (1),  ALBATROSS  sta.  2316, 
24°25'30"N,  81°47'45"W,  91  m;  USNM  74089  (1),  s  of  Key 
West;  USNM  117001  (1)  and  117002  (3),  Tortugas.  Gulf  coast 
of  Florida:  FMNH  64103  (1),  O-603;  FMNH  64104(1),  O-727- 
728;  FMNH  64109  (1),  O-732;  FMNH  64113  (1),  O-897; 
FMNH  64116  (2),  O-917;  FMNH  64119  (1),  O-936;  FMNH 
64122  (2),  O-945;  64126  (4),  O-1021;  FMNH  64129  (3),  O-1022; 
MCZ  45070  (1),  O-326;  USNM  134087  (1),  28°47'30"N, 
84°37'00"W.  Alabama:  FMNH  46744  (1),  O-265;  MCZ  45071 
(1),  0-2838;  USNM  159159  (1),  29°47'N,  87°17'W.  Louisiana: 
USNM  188793  (1),  SB- 175.  Yucatan:  USNM  188804  (1),  SB- 
404. 


Ogcocephalus  parvus  Longley  and  Hildebrand 

(Figure  28) 

Malthe  vespertilio  (non  Linnaeus):  GUNTHER  1880:7  [Pernam- 
buco(  =  Recife),  Brazil]. 

Ogcocephalus  pan-us  LONGLEY  AND  HILDEBRAND,  1940:283, 
fig.  28  [s  of  Tortugas,  Florida;  holotype  USNM  109313]; 
1941:311  and  314  [Tortugas,  Florida;  key];  ERDMAN 
1956:338  [Joyuda,  Puerto  Rico:  partial  description];  BRIGGS 
1958:301  [Florida,  compiled];  MOE  ET  AL.  1966:120  [Flori- 
da; list  of  museum  specimens];  BRADBURY  1967:417  [listed]; 
STARKS  1968:33  [Alligator  Reef,  Florida:  composition  of 
reef  fauna];  WALLS  1975:116  [N  Gulf  of  Mexico;  partial 
description]. 

Oncocephalus  parvus:  FOWLER  1945:336  [Key  West,  Florida; 
listed]:  1952:15  [Sombrero  Key  Light,  Florida:  description 
of  color]. 

Probable  reference: 

Ogcocephalus  vespertilio:  DAHLBERG  1975:45,  fig.  93  [coast 
of  Georgia:  partial  description;  range,  compiled]. 

DIAGNOSIS  AND  COMPARISONS. — An  Ogco- 
cephalus with  exceptionally  angular,  craggy 
body  contours  and  large,  prominent  bucklers. 
O.  parvus  differs  from  all  its  congeners  except 
cubifrons  in  having  a  very  small  mouth  relative 
to  head  depth  (width  of  mouth  into  head  depth 
2.0-2.8  in  parvus,  1.0-2.0  in  all  others  except 
cubifrons}.  O.  parvus  may  be  separated  from 
cubifrons  by  its  narrower  interorbital  space  (in- 
terorbital  width  into  head  depth  3.6-6.6  in  par- 
vus, 2.3-3.4  in  cubifrons).  It  may  be  distin- 
guished from  all  but  corniger,  declivirostris,  and 
pumilus  by  its  low  pectoral  fin  ray  count  (usually 
10-1 1  in  parvus,  12  or  more  in  all  others  except 
corniger,  declivirostris,  and  pumilus).  O.  par- 
vus is  distinguished  from  corniger  and  pumilus 
by  its  short  rostrum  (length  of  rostrum  into 
length  of  disk  margin  3.2-5.1  in  parvus,  1.7-3.0 
in  corniger  and  pumilus)  and  from  declivirostris 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


277 


FIGURE  28.     Ogcocephalus  parvus  Longley  and  Hildebrand.  Left  and  center:  FMNH  46742,  Gulf  of  Mexico,  75.0  mm  SL. 
Right:  FMNH  65957,  Guyana,  64.0  mm  SL. 


by  the  fleshy  ventral  pads  developed  on  the  dis- 
tal ends  of  the  pectoral  fin  rays  (absent  in  decli- 
virostris). 

DESCRIPTION. — Counts  and  measurements 
from  69  specimens,  27.3  to  84.7  mm  SL  (Table 
1).  Because  specimens  in  the  northern  part  of 
the  range  (coasts  of  southeastern  United  States 
and  eastern  Gulf  of  Mexico)  differed  in  pigment 
pattern  from  those  in  the  southern  part  (Carib- 
bean Sea  and  Atlantic  coast  of  South  America), 
the  two  groups  were  analyzed  separately  with 
the  expectation  that  samples  would  accumulate 
that  were  large  enough  to  differentiate  the 
groups  well.  However,  too  few  specimens  from 
the  southern  part  of  the  range  were  obtained,  so 
only  the  color  patterns  and  counts  are  treated 
separately  in  the  following  description.  Data  for 
body  proportions  were  combined  for  the  two 
groups. 

Counts.  Counts  given  in  Tables  2-6.  Counts 
for  parvus  the  lowest  found  in  genus.  Pectoral 
fin  ray  count  usually  10  in  northern  parvus  and 


11  in  southern  parvus;  range  10-12  for  species. 
Subopercular  lateral-line  count  usually  5  or  6; 
range  4-8.  Cheek  lateral-line  scale  count  usually 
8;  range  7-9.  Mean  count  for  lateral-line  scales 
17;  range  15-19.  Vertebral  count  modally  19; 
range  18-20  (all  vertebral  counts  are  from  ho- 
lotype  and  paratypes). 

Proportions.  Proportions  expressed  as  ratios 
given  as  mean  followed  by  range  in  parentheses. 
Relative  length  of  disk  margin  longest  for  the 
genus,  its  length  2.1(1.9-2.2)  in  SL.  Tail  mod- 
erately wide,  its  width  2.0(1.7-2.5)  in  length  of 
disk  margin.  Caudal  peduncle  3.1(2.5-4.0)  in 
head  depth.  Rostrum  variously  finger-  to  cone- 
shaped,  but  always  short,  its  length  4.2(3.2-5.1) 
in  length  of  disk  margin.  Aperture  of  illicial  cav- 
ity oval  or  subcircular.  Cranium  rising  steeply 
above  disk,  head  depth  1.8(1.6-2.0)  in  length  of 
disk  margin.  Width  of  cranium  3.2(2.6-3.6)  in 
length  of  disk  margin.  The  following  in  width  of 
cranium:  eye  1.4(1.1-1.6);  lateral  ethmoid  width 
1.7(1.4-1.9).  Interorbital  space  narrow,  con- 


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cave,  its  width  2.7(2.1-4.1)  in  width  of  cranium, 
4.8(3.6-6.6)  in  head  depth.  Mouth  small  and  nar- 
row, its  width  2.3(2.0-2.8)  in  head  depth, 
4.1(3.4-5.2)  in  length  of  disk  margin;  jaw  short, 
its  length  3.1(2.8-3.5)  in  head  depth.  Upper  lip 
fleshy,  of  an  even  width  for  its  entire  length; 
lower  lip  with  a  fleshy  median  lobe.  Dorsal  fin 
average  size,  its  length  3.4(2.4-4.3)  in  length  of 
disk  margin;  anal  fin  short,  its  length  3.1(2.4- 
3.6)  in  length  of  disk  margin. 

Integument.  Dermal  cirri  usually  sparse  or 
absent,  but  a  few  examples  have  cirri  well  de- 
veloped over  dorsal  surface  of  body.  Most  spec- 
imens from  southern  population  have  subter- 
minal  rostral  cirri.  Large  bucklers  arranged  as 
in  generic  description.  Dorsal  fin  devoid  of  tu- 
bercles; anal  fin  with  only  a  few  tiny  ones  at 
base.  Pectorals  and  pelvics  with  tiny  tubercles 
covering  basal  third  or  fourth  of  their  length. 
Caudal  as  in  generic  description.  Skin  covering 
eyeballs  bearing  tiny  tubercles  with  rim  around 
cornea  beaded  by  a  row  of  small  bucklers.  Pec- 
toral membranes  thick,  opaque;  ventral  surfaces 
of  ray  tips  with  well-developed  fleshy  pads. 

Color  in  preservative.  Northern  examples: 
ground  color  of  dorsal  surface  of  body  pale  tan 
to  medium  brown.  Markings  variable  but  irreg- 
ular or  rounded  dark  brown  spots,  with  or  with- 
out whitish  margins,  clustered  on  shoulders, 
face,  in  axillae,  and  sometimes  along  lateral 
sides  of  tail.  Occasionally  some  of  these  mark- 
ings are  rings  instead  of  spots.  Dark  splotches, 
sometimes  indistinct,  may  occur  elsewhere  on 
dorsal  surface.  A  few  specimens  with  minute 
dark  spots  scattered  over  dorsal  surface  in  ad- 
dition to  other  markings.  Ventral  surface  of 
body  uniformly  creamy  pale,  sometimes  becom- 
ing dusky  on  anterior  part  of  head,  particularly 
on  chin,  in  which  case  two  distinct  white  patches 
may  occur,  one  on  either  side  of  mandible.  Pec- 
torals pale  to  nearly  white  with  a  broad  black 
margin  extending  through  all  but  the  three  short- 
est rays;  ray  tips  white,  ventral  pads  white.  Pel- 
vics and  anal  same  pale  ground  color  as  ventral 
surface  of  body;  anal  often  with  a  dusky  or  dark 
spot  distally.  Dorsal  colorless  or  a  little  dusky 
basally.  Caudal  colorless  in  about  half  the  spec- 
imens at  hand,  otherwise  as  in  generic  descrip- 
tion. Iris  black  or  golden,  or  golden  with  dark 
spots. 

Southern  examples:  ground  color  of  dorsal 
surface  of  body  usually  a  much  darker  brown 


than  in  northern  specimens  (this  difference  does 
not  show  well  in  Fig.  28  because  photographed 
specimens  were  chosen  to  show  pigment  mark- 
ings contrasted  as  well  as  possible  with  ground 
color;  fish  on  right  in  Fig.  28  is  relatively  pale). 
Minute  but  very  distinct  black  dots  on  lateral 
portions  of  disk  and  on  top  of  cranium,  continu- 
ing posteriorly  along  dorsal  surface  of  tail. 
Large,  sharply  defined  brown  or  black  spots  and 
rings  on  shoulders  and  face  and  also  in  axillae, 
extending  from  these  on  to  sides  of  tail.  A  few 
specimens  with  an  absolutely  pigmentless  white 
patch  a  little  larger  than  top  of  cranium  in  area 
and  located  in  middorsal  region  just  posterior  to 
cranium.  Ventral  surface  of  body,  fins,  and  eyes 
as  in  northern  examples. 

Color  in  life.  From  an  aquarium  specimen 
newly  captured  in  Gulf  of  Mexico  off  Mobile, 
Alabama,  kindly  shown  me  by  Dr.  Robert  L. 
Shipp  and  his  students  at  the  University  of 
South  Alabama:  ground  color  of  face  and  dorsal 
surface  of  body  dark  brown,  the  large  irregularly 
shaped  spots  on  shoulders  and  axillae  lavender 
separated  by  cream-colored  reticula  in  approx- 
imately the  pattern  seen  on  the  right  in  Figure 
28;  smaller  pepperlike  spots  on  tail  and  disk  seen 
in  figure  are  black  in  living  specimen.  Down  face 
on  either  side  from  eye  to  angle  of  jaw  a  cascade 
of  orange  spots  having  dark  brown  rims  and  sep- 
arated by  white-colored  reticula.  Lips  orange. 
Iris  gold  flecked  with  orange.  Pectorals  a  trans- 
lucent orange  with  broad  black  margins.  Caudal 
dark  brown  basally,  white  in  middle  third,  or- 
ange on  distal  third. 

Several  Kodachrome  transparencies  made 
from  specimens  immediately  after  capture  have 
been  available.  From  specimens  from  off  the 
southeastern  United  States,  the  following  notes 
are  offered:  ground  color  of  dorsal  surface  of 
body  bright  tan  suffused  with  red-orange  in  large 
patches;  dark  spots  arranged  as  described  for 
preserved  specimens.  Dorsal  fin  red-orange. 
Pectorals  with  a  red-orange  stripe  just  proximal 
to  black  margin;  ray  tips  white. 

From  a  specimen  newly  captured  from  off 
coast  of  Guyana:  dorsal  surface  of  body  very 
dark,  the  markings  difficult  to  discern  except 
those  on  shoulders  (the  usual  cluster  of  irregular 
spots).  Skin  surrounding  gill  pores  tan.  Tips  of 
many  bucklers  faintly  red-orange.  Pectorals  with 
red-orange  stripe  just  proximal  to  black  margins; 
ray  tips  white.  Ventral  surface  of  body,  includ- 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


279 


ing  pectoral  peduncles  and  bases  of  pectoral 
fins,  entirely  bright  red-orange.  Anal  fin  red-or- 
ange with  white  stripe  through  second  quarter. 
Lips  red-orange,  corresponding  to  an  observa- 
tion by  Erdman  (1956)  of  a  specimen  from  Puer- 
to Rico  with  "bright  red  lips."  A  rectangularly 
shaped  white  patch  on  either  side  of  mandible 
and  an  oblong  white  patch  in  median  line  about 
halfway  between  chin  and  bases  of  pel  vies. 

DISTRIBUTION  (Fig.  25). — Known  from  the 
Atlantic  coast  of  the  United  States  from  Cape 
Hatteras  south  to  the  eastern  Gulf  of  Mexico, 
the  Caribbean  Sea,  and  the  coast  of  South 
America  to  Recife,  Brazil.  Bathy metric  range: 
29-126  m. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  For  data  from  early  OREGON  stations  (= 
O),  refer  to  Springer  and  Bullis  (1956);  for  data  from  later 
OREGON  stations  as  well  as  COMBAT  (=C)  and  SILVER  BAY 
(=SB)  stations,  refer  to  Bullis  and  Thompson  (1965). 
Northern  examples 

North  Carolina:  CAS  23922  (1),  SB-2927;  CAS  23924  (1), 
SB-3339;  FMNH  64336  (1),  C-384;  UF  24186  (1),  C-385.  South 
Carolina:  MCZ  45082  (1),  C-166.  Georgia:  FMNH  66383  (9), 
BOWERS  sta.  32,  30°14'N,  80°16'W,  73  m;  MCZ  45083  (1), 
PELICAN  sta.  178-15,  31°20'N,  80°17' W,  38  m;  UMML  1 14  (1), 
between  Jacksonville,  Florida,  and  Brunswick,  Georgia.  At- 
lantic coast  of  Florida:  ANSP  103631  (1),  SB-2721.  Florida 
Straits:  CAS-SU  62118  (2),  C-457;  UF  24187  (2),  SB-2363;  UF 
24188  (1),  SB-2382.  Florida  Keys:  FMNH  64125  (6),  O-1020; 
USNM  109313  (holotype)  and  USNM  109314  (12  paratypes), 
Tortugas,  134-201  m.  Gulf  coast  of  Florida:  FMNH  46742  (1), 
29°04'N,  84°23.5'W,  37  m;  FMNH  46743  (2),  O-35;  FMNH 
64105  (1),  O-729-730;  FMNH  64108  (2),  O-732;  FMNH  64112 
(1),  and  FMNH  64114  (2),  O-897;  FMNH  64115  (1),  O-916; 
FMNH  64117  (3),  O-917;  FMNH  64118  (2),  O-936;  FMNH 
64127  (3)  and  FMNH  64128  (3),  O-1021;  FMNH  64130  (1), 
O-1022;  GCRL  (1),  O-35;  MCZ  45084  (1),  O-1024;  USNM 
188795  (1),  PELICAN  sta.  153-3,  29°24'N,  85°54'W,  37  m.  Al- 
abama: USNM  188778  (1),  PELICAN  sta.  137-2,  29°36'N, 
87°29'W,  66  m.  Yucatan:  CAS  23923  (1),  SB-438;  FMNH 
46741  (1),  O-222. 
Southern  examples 

Puerto  Rico:  USNM  164504  (1),  Joyuda.  Honduras:  ANSP 
103630  (1),  O-1874.  Guyana:  CAS-SU  62121  (1),  O-2000; 
FMNH  64892  (2),  O-2247;  FMNH  64895  (3),  O-2261;  65954 
(1),  O-2245;  FMNH  65955  (1),  O-2232;  FMNH  65956  (1),  O- 
2257;  FMNH  65957  (4),  O-2000;  FMNH  65958  (1),  1999;  MCZ 
48081  (1),  O-2262;  USNM  188768  (2),  O-2000.  Brazil:  BMNH 
79.5.14.527  (1),  CHALLENGER  sta.  122,  off  Pernambuco  ^Re- 
cife). 

Ogcocephalus  darwini  Hubbs 

(Figure  29  [left]) 

Ogcocephalus  darwini  HUBBS,  1958:161  [Isla  Isabella,  Gala- 
pagos Islands;  holotype  SIO  H51-214;  photographs];  BRAD- 
BURY 1967:417  [listed];  MCALLISTER  1968:161  [Tagus  Cove, 
Albemarle  (  =  Isabella)  I.,  Galapagos  Islands;  branchioste- 
gals  described]. 


DIAGNOSIS  AND  COMPARISONS. — An  Ogco- 
cephalus distinguished  by  the  unique  character 
of  its  smooth  integument,  likened  to  shagreen 
by  Hubbs  in  the  original  description  (1958);  the 
bucklers  are  obscured  by  a  covering  of  fine  spi- 
nules  in  contrast  to  the  prominent,  coarse  buck- 
lers of  other  species.  O.  darwini  is  also  distin- 
guished from  all  its  congeners  except  the  single 
other  eastern  Pacific  species,  O.  porrectus,  by 
a  pair  of  solid  dark  stripes  on  the  dorsal  surface 
of  the  disk  which  continue  on  to  the  lateral  sides 
of  the  tail;  all  Atlantic  species  of  Ogcocephalus 
either  are  spotted  on  the  dorsal  surface  of  the 
body  or  else  lack  markings  altogether.  Differ- 
ences between  darwini  and  porrectus  include  a 
shorter  disk  margin  in  darwini  (Fig.  9)  and  a 
higher  modal  pectoral  fin  ray  count  (15  in  dar- 
wini, 14  in  porrectus;  Table  3). 

DESCRIPTION. — A  detailed  description  and 
comparison  with  the  two  other  eastern  Pacific 
ogcocephalines  is  given  in  the  original  descrip- 
tion of  Ogcocephalus  darwini  (Hubbs  1958),  but 
in  order  to  keep  the  presentations  of  species  in 
this  revision  strictly  parallel  for  ease  of  refer- 
ence, a  description  is  given  again  below,  includ- 
ing some  new  observations.  Counts  and  mea- 
surements from  30  specimens  35.1  to  166.1  mm 
SL  (Table  1). 

Counts.  Counts  given  in  Tables  2-6.  O.  dar- 
wini has  the  highest  mean  for  pectoral  fin  ray 
count  of  any  in  genus;  range  14-15  (but  Hubbs 
(1958)  gives  14-16;  however,  his  sample  includ- 
ed three  specimens  I  have  not  seen).  O.  darwini 
most  often  has  6-7  subopercular  lateral-line 
scales  but  shows  considerable  variation  in 
count;  range  4-9.  Cheek  lateral-line  count  usu- 
ally 8,  as  for  most  species  in  genus;  range  6-9. 
Range  for  lateral-line  scale  count  19-30,  which 
is  about  center  of  distribution  for  genus.  Modal 
vertebral  count  19;  range  19-20. 

Proportions.  Proportions  expressed  as  ratios 
given  as  mean  followed  by  range  in  parentheses. 

Disk  margin  shorter  on  the  average  than  in 
any  other  species  of  Ogcocephalus,  its  length 
2.6(2.3-2.9)  in  SL.  Tail  thin,  its  width  2.0(1.7- 
2.4)  in  length  of  disk  margin,  tapering  evenly  to 
caudal  fin;  caudal  peduncle  of  moderate  depth, 
2.7(2.4-3.1)  in  head  depth.  Rostrum  thick  and 
blunt,  moderately  long,  its  length  3.1(2.8-3.6)  in 
disk  margin.  Aperture  of  illicial  cavity  subtrian- 
gular,  higher  than  wide  or  the  two  dimensions 
about  equal.  Cranium,  when  viewed  from  front, 


280 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  7 


FIGURE  29.     Ogcocephalus  darwini  Hubbs,  left,  top  and  bottom,  paratype  128.5  mm  SL,  CAS-SU  17112.  Ogcocephalus 
porrectus  (Garman),  right,  top  and  bottom,  paratype  108.0  mm  SL,  MCZ  28733. 


rising  steeply  above  disk,  head  depth  1.8(1.6- 
2.1)  in  length  of  disk  margin.  Width  of  cranium 
3.2(2.9-3.5)  in  length  of  disk  margin.  The  follow- 
ing in  width  of  cranium:  eye  1.3(1.3-1.5),  lateral 
ethmoid  width  1.6(1.4-1.8).  Interorbital  space 
wide,  concave,  the  width  1.8(1.6-2.1)  in  width 
of  cranium,  3.2(2.9-3.6)  in  head  depth.  Mouth 
average  in  width,  the  width  1.7(1.4-2.0)  in  head 
depth,  3.1(2.7-3.4)  in  length  of  disk  margin; 


length  of  jaw  2.4(2.0-2.6)  in  head  depth.  Upper 
lip  moderately  fleshy,  of  an  even  width  for  its 
entire  length;  lower  lip  fleshy,  no  median  lobe. 
Dorsal  fin  average  size,  its  length  3.8(3.0-4.3)  in 
length  of  disk  margin;  anal  fin  comparatively 
long,  2.4(2.1-2.6)  in  length  of  disk  margin. 

Integument.  Dermal  cirri  present  or  absent. 
Large  bucklers  present  but  inconspicuous  be- 
cause of  their  covering  of  skin  embedded  with 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


281 


E 

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cu 

o 

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, 

I 

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50 


60 


70 


80 


130 


140 


150 


160 


90  100  110  120 

Standard  length  in  mm 

FIGURE  30.     Comparison  between  Ogcocephalus  darwini  and  O.  porrectus  to  show  that  length  of  disk  margin  is  greatest  in 
O.  porrectus  relative  to  standard  length. 


fine  spinules  (except  small  specimens  35  mm  SL 
or  less  have  perfectly  visible  bucklers  not  yet 
covered  by  the  fine-grained,  shagreenlike  integ- 
ument); bucklers  arranged  as  in  generic  descrip- 
tion. All  fins  with  fine  tubercles  at  their  bases 
and  extending  out  variable  distances  along  fin 
rays.  Caudal  as  in  generic  description.  Skin  cov- 
ering eyeballs  bearing  tiny,  densely  set  tuber- 
cles, with  rim  around  cornea  beaded  by  a  row 
of  somewhat  larger  ones.  Pectoral  membranes 
thick,  opaque;  ventral  surfaces  of  ray  tips  with 
well-developed  fleshy  pads. 

Color  in  preservative.  Ground  color  of  dorsal 
surface  tan  to  brown,  the  conspicuous  markings 
a  pair  of  dark  stripes,  one  on  each  side,  origi- 
nating behind  eyes  and  extending  posteriorly 
over  disk  and  on  to  lateral  walls  of  tail  where 
they  are  sometimes  interrupted  to  form  a  series 
of  blotches.  Top  of  head  dark,  from  whence  a 
median  dusky  stripe  extends  posteriorly,  be- 
coming wider  and  more  intense  around  dorsal 
fin.  Face  also  dark,  marked  by  a  narrow  light 
stripe  on  either  side  extending  obliquely  from 
eye  to  lip  just  anterior  to  corner  of  mouth,  a 
pattern  very  similar  to  that  observed  in  Ogco- 
cephalus porrectus.  Ventral  surface  uniformly 
pale  except  for  chin,  which  is  sometimes  dusky; 
the  pale  shade  extends  up  sides  of  tail  to  the 
lateral  line  or  even  a  little  above.  Rostrum  the 
same  dark  shade  as  face  and  head.  Iris  black  or 
gold,  or  black  with  golden  spots.  Dorsal  surfaces 
of  pectorals  pale  basally,  grading  to  black  dis- 
tally,  the  ventral  pads  near  tips  of  rays  pale  or 


white.  Pelvics  and  anal  the  same  ground  color 
as  ventral  surface  of  body,  anal  sometimes 
dusky  or  black  on  distal  third.  Dorsal  fin  dusky 
to  dark,  sometimes  blotched.  Caudal  fin  pale 
basally  with  distal  third  dark  except  in  one  spec- 
imen, which  displays  color  pattern  described  for 
genus. 

Color  in  life.  Hubbs  ( 1958)  states  for  one  spec- 
imen which  retained  some  color,  "upper  parts, 
purplish  gray;  rather  blue-gray  on  tubercles  and 
in  an  irregular  blotch  near  each  side  of  the  disk; 
the  two  dark  streaks,  reddish  brown;  under- 
parts,  bright  rose-red,  becoming  white  or  whit- 
ish on  the  lower  (but  not  the  upper)  surface  of 
the  pelvics,  on  the  outer  tip  of  the  anal,  and, 
weakly,  on  the  lower  border  of  the  caudal,  also 
on  the  esca  (but  not  the  red  stem)  of  the  illicium; 
pectoral  rays  pink-gray,  encroached  by  the  wid- 
ening, blackish  interradial  streaks;  the  fin  be- 
coming almost  solidly  blackish  inside  the  narrow 
red  outer  border." 

I  observed  two  live  specimens  in  August  1968 
which  had  been  freshly  taken  from  coral  rubble 
in  25  m  of  water  off  Isla  Isabela  by  pipe  dredge. 
In  these  the  dorsal  surfaces  of  disk  and  tail  me- 
dium brown,  the  two  longitudinal  stripes  dark 
cocoa  brown.  Ventral  surfaces  of  body  creamy 
white,  as  were  esca  and  tops  of  eyeballs,  all  con- 
trasting sharply  with  brown  head  and  brown  ros- 
trum. Lips  cherry-red;  striking  white  blotches 
along  sides  of  disk  posterior  to  mouth.  Pelvics 
creamy  white  tipped  with  brown  on  dorsal  sur- 
faces. Pectorals  brown  on  dorsal  surfaces,  grad- 


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ing  to  rich  dark  shade  distally,  creamy  white  on 
ventral  surfaces. 

DISTRIBUTION. — Galapagos  Islands.  Bathy- 
metric  range:  3.5-73.5  m. 

MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  All  material  from  the  Galapagos  Islands. 
Isla  Isabels:  SIO  H51-214  (holotype)  and  SIO  54-175  (1),  Cale- 
ta  Tagus;  SIO  H50-18  (1),  Punto  Moreno,  shallow  water  near 
shore;  SIO  55-16  (1),  Punto  Moreno,  under  9  m;  SIO  H50-132 
(1),  Bahia  Elizabeth,  5.5  m;  SIO  54-199  (1)  and  SIO  58-116 
(1),  Bahia  Elizabeth;  SIO  57-20  (1),  Bahia  de  Banks,  approx. 
00°01'S,  91°29'W;  CAS-SU  14977  (1),  Bahia  de  Banks,  under 
77  m;  SIO  H51-51  (1),  just  outside  Caleta  Webb;  SIO  57-111 
(1),  Caleta  Webb;  CAS  39904  (13),  TE  VEGA  cr.  17,  sta.  91, 
00°15'22"S,  91°22'26"W,  Canal  Bolivar;  SIO  56-60  (1),  4.6-5.6 
m;  SIO  58-39  (1),  w  side  I.  Isabela  or  E  side  I.  Fernandina. 
Isla  Fernandina:  SIO  H53-196  (1),  Punt  a  Mangle.  Isla  Santa 
Cruz:  USC  (1),  Allan  Hancock  Pacific  Exped.  1935,  sta.  345- 
35,  00°24'50"S,  90021'40"W,  55  m. 

Other  material:  SIO  57-166  (1),  exact  locality  unknown; 
CAS-SU  17112  (1),  exact  locality  unknown;  CAS-SU  46654 
(1),  from  stomach  of  shark  (Gyropleurodus  quoyi)  taken  in 
Caleta  Tagus. 

Ogcocephalus  porrectus  (Garman) 
(Figure  29  [right]) 

Oncocephalus  porrectus  GARMAN,  1899:86  [5°32'45"N, 
86°54'30"W;  lectotype  MCZ  28733]. 

Ogcocephalus  porrectus:  HUBBS  1958:161  [redescription  of 
type-series;  photographs;  selection  of  lectotype];  BRAD- 
BURY 1967:417  [listed]. 

DIAGNOSIS  AND  COMPARISONS. — One  of  only 
two  species  of  Ogcocephalus  known  from  the 
eastern  Pacific  Ocean  (the  other  is  O.  darwini). 
O.  porrectus  and  O.  darwini  are  morphologi- 
cally distinguishable  by  the  nature  of  the  squa- 
mation,  shagreenlike  and  relatively  smooth  in 
darwini  but  rough  with  prominent  spiny  buck- 
lers in  porrectus.  Other  differences  include  the 
relatively  longer  disk  margin  in  porrectus  (Fig. 
9)  and  lower  modal  number  of  pectoral  rays  (14 
in  porrectus,  15  in  darwini,  Table  3). 

Although  a  smaller  species  than  nasutus  (Ta- 
ble 1),  porrectus  most  resembles  nasutus  in 
body  proportions  and  quality  of  the  squamation. 
However,  the  color  pattern  in  porrectus,  con- 
sisting of  a  longitudinal  stripe  on  each  side  of 
the  body,  is  unknown  in  nasutus  or  any  other 
Atlantic  species.  The  relationship  in  color  pat- 
tern between  Atlantic  and  Pacific  species  in  this: 
the  clusters  of  spots  or  reticulations  where  found 
in  tracts  in  Atlantic  species  are  represented  in 
porrectus  (and  in  darwini)  by  solid  stripes. 

DESCRIPTION. — Counts  and  measurements 
from  35  specimens  25.6  to  138.5  mm  SL  (Table 
1). 

Counts.  Pectoral  fin  ray  count  relatively  high 


with  modal  number  14  in  range  of  10  to  15  for 
genus  (Table  3).  Lateral-line  scale  counts  below 
average,  however,  with  lateral-line  count  always 
under  30  (Table  4)  and  subopercular  lateral-line 
count  modally  6  (Table  6).  Cheek  lateral-line 
scale  count  usually  8  as  in  most  species  in  the 
genus  (Table  5)  and  vertebral  count  usually  19 
(Table  2). 

Proportions.  Proportions  expressed  as  ratios 
given  as  mean  followed  by  range  in  parentheses. 
Disk  outline  subtriangular,  length  of  disk  margin 
2.2(2.1-2 .4)  in  SL.  Tail  thin  to  moderately  wide, 
its  width  2.0(1.6-2.4)  in  length  of  disk  margin, 
tapering  evenly  to  caudal  fin;  caudal  peduncle 
of  average  thickness,  its  depth  3.0(2.6-3.4)  in 
head  depth.  Rostrum  moderately  long  with  a 
thick  base,  its  length  3.0(2.8-3.4)  in  length  of 
disk  margin;  seen  from  side,  rostrum  arches 
slightly  with  its  distal  end  pointing  downwards 
except  that  terminal  buckler  is  turned  abruptly 
upwards  (Fig.  29).  Aperture  of  illicial  cavity  sub- 
triangular  in  outline,  higher  than  wide.  Cranium, 
when  viewed  from  front,  rising  steeply  above 
disk;  head  depth  2.0(1.9-2.2)  in  length  of  disk 
margin.  Width  of  cranium  3.2(2.3-3.9)  in  length 
of  disk  margin.  The  following  in  width  of  cra- 
nium: eye  1.6(1.3-2.0),  lateral  ethmoid  width 
1.7(1.5-1.9).  Interorbital  space  flat,  not  convex 
as  in  vespertilio,  of  average  width,  the  width 
2.1(1.8-2.5)  in  width  of  cranium,  3.6(3.0-4.0)  in 
head  depth.  Mouth  moderate,  its  width  1.6(1.4- 
1.8)  in  head  depth,  3.2(2.8-3.6)  in  length  of  disk 
margin;  length  of  jaw  2.3(2.1-2.5)  in  head  depth. 
Upper  lip  moderately  fleshy,  of  an  even  width 
for  its  entire  length;  lower  lip  fleshy,  thickened 
medially.  Dorsal  fin  small,  its  length  4.4(3.4—4.8) 
in  length  of  disk  margin;  anal  fin  relatively  long, 
its  length  2.6(2.4-3.0)  in  length  of  disk  margin. 

Integument.  Dermal  cirri  inconspicuous,  usu- 
ally present  on  disk  margin,  chin,  lateral  sides 
of  tail  near  lateral  line,  and  on  sides  of  large 
bucklers  on  dorsal  surface  of  body.  Large  buck- 
lers prominent,  with  coarse  spines  bristling  from 
apex  of  each;  bucklers  arranged  as  in  generic 
description.  All  fins  except  dorsal  with  fine  tu- 
bercles running  out  for  l/3  to  %  length  of  fin  rays; 
dorsal  fin  with  none  or  only  a  few  prickles  on 
anterior  edge  of  first  ray.  Caudal  as  in  generic 
description.  Skin  covering  eyeballs  bearing  fine 
tubercles  with  rim  around  cornea  beaded  by  a 
row  of  small  but  prominent  bucklers.  Pectoral 
membranes  thick,  opaque;  ventral  surfaces  of 
ray  tips  with  well-developed  fleshy  pads. 

Color  in  preservative.  Freshest  material  has 


BRADBURY:  FISH  GENUS  OGCOCEPHALUS 


283 


dorsal  surface  of  body  uniformly  brown  except 
for  darker  brown  longitudinal  stripes,  one  on 
each  side  as  in  darwini,  beginning  anteriorly  on 
dorsal  surface  of  disk  behind  eyes  as  an  elongate 
"shoulder"  blotch  which  then  narrows  poste- 
riorly and  trails  back  along  sides  of  tail  where 
sometimes  interrupted  in  one  or  two  places. 
Carman's  four  specimens  faded  (Garman  1899), 
but  holotype  (Hubbs  1958)  shows  the  longitu- 
dinal markings,  and  original  description  refers 
to  markings. 

Markings  on  faces  of  fresh  specimens  also  re- 
semble markings  in  darwini;  suborbital  space 
(between  eye  and  mouth)  dark  except  for  a 
markedly  pale  stripe  descending  from  eye  to 
corner  of  mouth.  Lips  the  same  creamy- white 
shade  as  ventral  surface  of  body;  pelvics  and 
ventral  surfaces  of  pectorals  also  creamy  white. 
Dorsal  surface  of  pectorals  dusky,  grading  dis- 
tally  to  black  tips.  Anal  often  tipped  with  black, 
especially  in  small  specimens.  Dorsal  dusky, 
occasionally  blotched  with  dark  pigment.  Iris 
golden  with  very  dark  spots  in  a  ring  around  the 
eccentrically  shaped  pupil  (Garman  described 
the  iris  as  having  "radiating  bars  of  brown"). 
Caudal  as  in  generic  description. 

Color  in  life.  Two  35-mm  color  slides  provided 
by  Dr.  Robert  Lea  show  that,  as  in  other  species 
of  Ogcocephalus,  O.  porrectus  has  considerable 
bright-reddish  coloring.  One  slide  shows  entire 
ventral  surface  of  body  to  be  orange-red  except 
for  ventral  surfaces  of  pelvic  and  anal  fins, 
which  are  whitish.  The  other  slide,  a  dorsal  view 
of  body,  shows  skin  around  gill  openings  to  be 
reddish  and  fin  rays  nearest  sides  of  pectoral  and 
caudal  fins  to  have  their  tips  scarlet. 

DISTRIBUTION. — Vicinity  of  Cocos  Island 
from  depths  of  88-146  m.  Hubbs  (1958)  erred  in 
stating  that  the  material  described  by  Garman 
(1899)  from  ALBATROSS  station  3368  was  from 
"south  of  the  Gulf  of  Panama,  in  the  vicinity  of 
Cabo  Corrientes,  Colombia."  The  coordinates 
for  this  station  (5°32'45"N  and  86°54'30"W,  cor 
rectly  quoted  by  Prof.  Hubbs)  designate  a  lo- 
cality near  Cocos  Island,  not  the  coast  of  Co- 
lombia. But  this  small  error  may  foreshadow 
things  to  come.  A  specimen  from  Peru,  USNM 
200363  (data  given  below)  agrees  fairly  well  with 
O.  porrectus  in  squamation,  color  pattern,  and 
morphometric  characters.  Whether  a  population 
of  batfishes  that  properly  may  be  assigned  to  O. 
porrectus  occurs  along  the  coast  of  western 
South  America  is  a  question  that  awaits  future 
work. 


MATERIAL  EXAMINED. — Numbers  in  parentheses  are  num- 
bers of  specimens.  Cocos  Island:  MCZ  28733  (lectotype)  and 
MCZ  41594  (3  paralectotypes)  both  from  ALBATROSS  sta. 
3368,  5°32'45"N,  86°54'30"W,  121  m.  The  following  from  R/V 
SEARCHER  cruise  72-4:  LACM  32263  (23),  5°33'32"N, 
87°04'44"W,  110  m;  LACM  32264  (6),  5°33'30"N,  87°05'50"W, 
137-146  m;  LACM  32268  (1),  5°31'10"N,  87°01'58"W,  88-91 
m;  LACM  32269  (1),  5°28'30"N,  87°04'00"W,  119-121  m. 

Peru:  USNM  200363  (1),  Caleta  Cruz  (Tumbes),  37-55  m. 


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the  Oregon  in  the  Gulf  of  Mexico.  U.S.  Fish  Wildl.  Serv. 

Spec.  Sci.  Rep.  Fish.  196:1-134. 
SPRINGER,  V.  G.,  AND  A.  J.  MC£RLEAN.    1962.  Seasonality 

of  fishes  on  a  south  Florida  shore.  Bull.  Mar.  Sci.  Gulf 

Caribb.  12(1):39-60. 
-  ,  AND  K.  D.  WOODBURN.    1960.   An  ecological  study 

of  the  fishes  of  the  Tampa  Bay  area.  Florida  State  Bd  .  Cons. 

Mar.  Lab.,  Prof.  Pap.  Ser.  1.  104  p. 
STARK,  W.  A.,  II.    1968.    A  list  of  fishes  of  Alligator  Reef, 

Florida,  with  comments  on  the  nature  of  the  Florida  reef 

fish  fauna.  Undersea  Biol.  1(1):4-40. 
STORER,  D.  H.   1846.  A  synopsis  of  the  fishes  of  North  Amer- 

ica. Mem.  Am.  Acad.  Arts  Sci.  Boston  2(7):44-298. 
STOREY,  M.,  AND  E.  W.  GUDGER.   1936.  Mortality  of  fishes 

due  to  cold  at  Sanibel  Island,  Florida,  1886-1936.  Ecology 

17(4):  640-648. 
UHLER,  P.  R.,  AND  O.  LUGGER.   1876.  List  of  fishes  of  Mary- 

land. Rep.  Comm.  Fish  Maryland  1876:67-176. 
WALLS,  J.  G.    1975.   Fishes  of  the  northern  Gulf  of  Mexico. 

T.F.H.  Publ.,  New  Jersey.  432  p. 
WOODS,  L.  P.    1942.    Rare  fishes  from  the  coast  of  Texas. 

Copeia  1942(3):  191. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  8,  pp.  287-293,  5  figs.  July  2,  1980 


XENAPLOACTIS,  A  NEW  GENUS  FOR  PROSOPODASYS  ASPERRIMUS 

GUNTHER  (PISCES:  APLOACTINIDAE),  WITH 

DESCRIPTIONS  OF  TWO  NEW  SPECIES 

By 
Stuart  G.  Poss 

Division  of  Biological  Sciences  and  Museum  of  Zoology, 
University  of  Michigan,  Ann  Arbor,  Michigan  48109 

and 
William  N.  Eschmeyer 

California  Academy  of  Sciences,  Golden  Gate  Park, 
San  Francisco,  California  94118 


ABSTRACT.  A  new  genus,  Xenaploactis,  is  created  for  Prosopodasys  asperrimus  Giinther,  1860,  which  is  re- 
described.  \.  anopta  from  Luzon  Island  in  the  Philippines  and  A .  cautes  from  the  Andaman  Sea  and  the  Gulf 
of  Thailand  are  described  as  new.  These  species  exhibit  differences  in  the  configuration  of  ridges  between  the 
eyes,  body  depth,  head  pores,  and  other  features.  Species  of  Xenaploactis  differ  from  those  of  other  genera  of 
the  Aploactinidae  by  the  presence  of  several  features  in  combination:  3  anterior  dorsal  fin  spines  forming  a 
separate  fin,  rather  sharp  head  spines,  a  markedly  upturned  mouth,  and  a  body  densely  covered  with  modified 
pointed  scales. 

INTRODUCTION  view.  More  recent  examination  of  the  holotype 

Giinther  (1860)  described  Prosopodasys  as-  confirms  this  suspicion. 

perrimus,  assigning  it  to  a  genus  otherwise  com-  Species  assigned  to  Prosopodasys,  a  name 

posed  of  tetrarogid  scorpaenoids.  Because  of  originally  proposed  by  Cantor  (1849:1026)  as  a 

this  and  because  the  species  remains  known  replacement  name  for  Apistus  Cuvier  and  Val- 

from  only  the  holotype,  Prosopodasys  asperri-  enciennes,  have  been  placed  into  a  number  of 

mus  has  not  been  generally  recognized  as  an  genera.  Prosopodasys  asperrimus  does  not  be- 

aploactinid  scorpaenoid.  long  in  any  of  these  nor  does  it  belong  in  any 

Giinther  described  the  head  and  body  as  existing  genus  of  aploactinid. 

"covered  with  small  prickles,"  and  this  caused  Two  undescribed  and  closely  related  species 

Poss  and  Eschmeyer  (1978)  to  suspect  that  this  have  been  discovered  among  specimens  in  the 

species  was  an  aploactinid.  Other  features  men-  collections  of  the  National  Museum  of  Natural 

tioned  in  Giinther's  brief  description — absence  History  (USNM)  and  the  California  Academy  of 

of  palatine  teeth,  presence  of  one  spine  and  Sciences  (CAS).  Together  with  Prosopodasys 

three  soft  rays  in  the  pelvic  fins,  and  presence  asperrimus,   they  form  a  natural  cluster  quite 

of  flexible  anal  spines — tended  to  support  this  distinct  from  other  aploactinids.  Provided  below 

[287] 


288 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  8 


are  descriptions  of  these  species  and  a  new  ge- 
nus to  contain  them. 

METHODS 

The  methods  used  in  this  study  follow  those 
of  Eschmeyer  (1969)  as  modified  by  Poss  and 
Eschmeyer  (1978).  Spines  and  rays  are  difficult 
to  distinguish  and  median  fin-ray  counts  were 
checked  against  radiographs.  The  last  fin  ray  in 
the  dorsal  and  anal  fins  is  double,  borne  on  a 
single  pterygiophore,  and  counted  as  one  ray. 

Xenaploactis  new  genus 
TYPE-SPECIES. — Prosopodasys  asperrimus  Giinther,  1860. 

DIAGNOSIS. — Aploactinid  fishes  with  spinous 
dorsal  fin  in  2  parts;  the  first  3  dorsal  fin  spines 
inserting  on  cranium  and  forming  a  separate  fin, 
widely  separate  from  the  4th  spine  of  the  second 
part  of  the  spinous  dorsal  fin;  rather  pungent 
head  spines,  including  2  prominent  preorbital 
spines  and  strong  spine  on  lateral  face  of  third 
infraorbital  bone;  head  and  body  densely  cov- 
ered with  modified  scales;  mouth  strongly  up- 
turned. Maxillary  with  prominent  angular  point 
at  end  of  anteroventral  margin,  with  ridge  on 
lateral  face.  Dorsal  HI,  X,  8-9  (last  double)  or 
III,  XI,  8-9  (last  double).  Anal  I,  9-10  (last  dou- 
ble). Pelvic  I,  3.  Pectoral  13-14.  Vertebrae  27- 
28.  Branchiostegal  rays  6.  Four  dorsal  spines 
and  associated  pterygiophores  anterior  to  third 
neural  spine.  Upper  posterior  margin  of  oper- 
cular  bone  very  close  to  base  of  dorsal  fin.  Teeth 
on  jaws  and  vomer,  none  on  palatines. 

DESCRIPTION. — (See  also  species  descriptions 
below.)  Dorsal  fin  in  2  parts,  originating  on  cra- 
nium above  posterior  border  of  eye;  first  3 
spines  close  together,  widely  separate  from  4th 
spine,  which  originates  just  anterior  to  end  of 
head;  2nd  spine  longest,  all  spines  rather  weak, 
flexible,  without  cirri.  Dorsal  fin  membrane  of 
second  part  of  fin  notably  incised,  dorsal  spines 
free  from  fin  membrane  at  about  midlength;  de- 
gree of  fin  incision  decreases  posteriorly.  Pec- 
toral fin  rays  all  unbranched,  longest  5-7  from 
above,  reaching  just  past  anus. 

Head  covered  with  modified  scales,  scales  ab- 
sent in  interorbit  and  behind  eye.  Mouth  strong- 
ly upturned.  Movable  lachrymal  bone  (infraor- 
bital 1)  bladelike,  with  3  spines;  1st  small, 
directed  anteriorly,  followed  by  2  prominent 
sharp  spines,  2nd  spine  about  equal  in  length  to 
3rd,  directed  downward;  3rd  spine  directed 


down  and  slightly  back.  Third  infraorbital  with 
prominent  spine  on  ventral  margin  of  bone  pro- 
jecting ventrally  and  laterally;  lateral  face  of 
bone  with  prominent  ridge  and  2  blunt  spines. 
Infraorbital  (suborbital)  stay  appearing  as  a 
strong  ridged  bar.  One  postorbital  bone  (infraor- 
bital 4  or  5).  Interorbit  with  prominent  ridges. 
Nasal  bones  tubed,  without  spines.  Preopercle 
with  5  rather  strong  spines,  uppermost  largest, 
diminishing  in  size  ventrally.  Upper  arm  of  pre- 
opercle  forming  strong  ridge.  Opercle  with  2 
weak  ridges,  lower  ending  in  small  blunt  spine 
on  opercular  margin.  Opercular  flap  extending 
nearly  to  base  of  dorsal  fin  between  spines  4  and 
5.  Interopercle  forming  spinous  projection  on 
posterior  opercular  margin.  Posterior  dorsal 
border  of  cleithrum  ending  in  small,  poorly  de- 
fined blunt  spine.  Angular  bone  prominent, 
strongly  jutting  ventrally  with  mouth  closed. 
Maxillae  reaching  anterior  border  of  eye,  with 
ridged  dorsal  border,  ridge  near  ventral  border. 
Maxillary  cirrus  minute. 

Pelvic  fin  origin  slightly  in  advance  of  lower- 
most pectoral  fin  rays.  Pelvic  fin  membrane  not 
adnate  to  body.  Caudal  fin  rounded,  with  18  total 
fin-ray  elements  (9  upper  and  9  lower),  all  un- 
branched. Caudal  skeleton  with  parahypural,  1st 
and  2nd  hypurals  fused;  3rd  and  4th  hypurals 
fused;  5th  hypural  small,  autogenous;  1st  preural 
neural  spine  long;  2  epurals.  Seven  upper  and  7 
lower  fin-ray  elements  are  attached  to  the  hy- 
purals, 2  (procurrent)  rays  are  free  above  and 
below. 

ETYMOLOGY. — The  generic  name  Xenaploac- 
tis is  derived  from  the  Greek  xenos  (stranger)  + 
Aploactis  (a  related  genus).  Xenaploactis  is 
feminine. 

Key  to  the  Species  of  Xenaploactis 

la.  Interorbit  with  ridges  nearly  parallel  (Fig. 
4).  Body  depth  less  than  Vz  of  standard 
length.  Dorsal  fin  III,  IX,  8-9  (based  on 
limited  material;  some  variation  to  be  ex- 
pected) .___  X.  cautes  (Figs.  4  lower  and  5) 

Ib.  Interorbit  with  ridges  divergent  anterior- 
ly, convergent  over  middle  of  orbit,  di- 
vergent posteriorly  (Figs.  2  and  4  upper). 
Body  depth  equal  to  or  greater  than  Ys  of 
standard  length.  Dorsal  fin  III,  X,  8-9 
(based  on  limited  material;  some  variation 
to  be  expected) 2 

2a.  Second  infraorbital  bone  with   1   or  2 


POSS  &  ESCHMEYER:  NEW  APLOACTINID  GENUS  AND  SPECIES 


289 


FIGURE  1.    Lateral  view  of  holotype  of  Prosopodasys  asperrimus  (=Xenaploactis  asperrima)  (BMNH  1979.5.5: 1,  39.9  mm 
SL).  Specimen  formerly  dried,  somewhat  distorted. 


spines.  Pore  of  infraorbital  lateral  line  ca- 
nal at  second  infraorbital  bone  as  simple 
obscure  pore.  Body  depth  greater  than  l/3 
of  standard  length.  No  fingerlike  cirri 
above  uppermost  preopercular  lateral  line 
pores.  Dorsal  fin  III,  X,  9.  Anal  I,  10 
(based  on  limited  material;  some  variation 

to  be  expected) 

X.  asperrima  (Figs.  1  and  2) 

2b.  Second  infraorbital  bone  without  spines. 
Pore  of  infraorbital  lateral  line  canal  at 
second  infraorbital  bone  as  prominent 
elongate  slit.  Body  depth  equal  to  Vz  of 
standard  length.  Fingerlike  cirri  present 
above  uppermost  preopercular  lateral  line 
pores.  Dorsal  III,  X,  8.  Anal  I,  9  (based 
on  limited  material;  some  variation  to  be 

expected) 

X.  anopta  (Figs.  3  and  4  upper) 


Xenaploactis  asperrima  (Gunther) 
(Figures  1  and  2) 

Prosopodasys  asperrimus  GUNTHER,  1860:140-141  (original 
description;  type-locality  East  Indies). 

MATERIAL.— Holotype:  BMNH  1979.5.5:1  (39.9  mm  SL). 
East  Indies,  Sir  E.  Belcher,  no  other  data. 

COUNTS.— Dorsal  fin  III,  X,  9  (last  double). 
Anal  fin  I,  10  (last  double).  Pectoral  fin  13  (left), 
14  (right).  Pelvic  fin  1,  3.  Lateral  line  scales  10 
(left),  11  (right).  Vertebrae  27. 

DESCRIPTION. — (See  also  generic  diagnosis 
above.)  Body  notably  elevated  behind  head, 


body  depth  more  than  Vs  of  standard  length. 
Body  densely  covered  with  modified  scales 
which  form  spinous  points,  best  developed  on 
upper  back  behind  head.  Lateral  line  with  10-11 
tubed  scales,  each  with  2  small  laterally  pro- 
jecting spinules,  best  developed  anteriorly,  last 
scale  extending  over  base  of  caudal  fin.  Gill  rak- 
ers short,  difficult  to  count,  total  8-10;  3  on  up- 
per arch,  5-7  on  lower  arch.  No  modified  scales 
on  snout.  Lachrymal  bone  (infraorbital  1)  with 
2  large  spines,  first  notably  curved.  A  small 
spine  in  front  at  base  of  first  spine,  a  small  spine 
at  base  of  second  spine.  Second  infraorbital 
bone  with  a  small  double  or  single  spine,  with 
obscure  small  circular  lateral  line  pore.  Third 
infraorbital  bone  with  a  large  spine  directed  out 
and  down;  a  strong  ridge  attached  to  preopercle. 
Mouth  very  strongly  upturned,  nearly  vertical. 
Interorbit  with  prominent  ridges,  divergent  an- 
teriorly, convergent  over  middle  of  interorbit, 
divergent  posteriorly  (Fig.  2).  Postocular  spine 
appearing  as  a  sculptured  ridge,  weakly  con- 
nected to  supraorbital  ridge.  Parietal  spine 
lumplike.  Pterotic  spine  as  a  strong,  slightly 
curved  ridge.  Posttemporal  spine  well  ossified, 
sculptured,  appearing  as  a  ridged  lump,  followed 
by  small  bladelike  supracleithral  spine.  Dorsal 
posterior  border  of  cleithrum  appearing  as  a 
marked  ridge,  ending  in  a  blunt,  poorly  defined 
spine.  Preopercular  lateral  line  pores  simple,  no 
fingerlike  cirri  above  uppermost  pores.  Ventral 
surface  of  dentary  without  distinct  cirri. 
Color  in  life  unknown.  Color  of  head  and  body 


290 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  8 


FIGURE  2.     Dorsal  view  of  head  of  holotype  of  Prosopo- 
dasys  asperrimus  (=Xenaploactis  asperrima). 


in  preservative  light  brown,  fins  slightly  darker 
and  possibly  speckled  in  life. 

Measurements  in  millimeters  as  follows  (per- 
cent standard  length  in  parentheses):  standard 
length  39.9;  head  13.2  (34);  snout  3.5  (9);  orbit 
3.0  (7);  interorbital  width  2.2  (5);  jaw  6.0  (15); 
postorbit  7.0  (17);  body  depth  14.6  (37);  predor- 
sal  6.2  (15);  anal  fin  14.0  (35);  caudal  fin  9.5  (24); 
pectoral  fin  9. 2  (23);  pelvic  fin  4. 5  (11);  1st  dorsal 
spine  3. 2  (8);  2nd  dorsal  spine  4.1  (10);  3rd  dorsal 
spine  3.1  (8);  4th  dorsal  spine  1.8  (4);  5th  dorsal 


spine  2.7  (7);  penultimate  dorsal  spine  3.1  (8); 
last  dorsal  spine  3.5  (9);  anal  spine  2.1  (5);  least 
depth  of  caudal  peduncle  4.2  (10);  snout  to  base 
of  2nd  dorsal  spine  7.2  (18);  snout  to  base  of  3rd 
dorsal  spine  7.8  (19);  snout  to  base  of  4th  dorsal 
spine  1 1.5  (29);  snout  to  base  of  5th  dorsal  spine 
14.6  (37);  width  of  1st  dorsal  spine  at  midlength 
0.2  (1);  incision  of  dorsal  fin  membrane  at  4th 
dorsal  spine  (from  tip  to  membrane)  1.8  (4). 

DISTRIBUTION. — Known  only  from  the  holo- 
type from  the  "East  Indies." 

Xenaploactis  anopta,  new  species 

(Figures  3  and  4  upper) 

No  literature  applies  to  this  species. 

MATERIAL.— Holotype:  CAS  32633  (37.0  mm  SL).  Philip- 
pines, Luzon  I.,  Zambales,  4  km  w  of  Calguaguin  Cove,  64- 
81  m,  0835-0910  hrs,  J.  E.  Norton,  9  June  1966. 

COUNTS. — Dorsal  fin  III,  X,  8  (last  double). 
Anal  fin  I,  9  (last  double).  Pectoral  fin  13  (left), 
14  (right).  Pelvic  fin  I,  3.  Lateral  line  scales  10 
(left),  11  (right).  Vertebrae  27. 

DESCRIPTION. — (See  also  generic  description 
above.)  Body  somewhat  elevated  behind  head, 
body  depth  Vs  of  standard  length.  Body  densely 
covered  with  modified  scales  which  form  spi- 
nous  points,  best  developed  anteriorly;  lateral 
line  with  10-11  tubes,  each  with  2  small,  later- 
ally projecting  spinules  which  are  best  devel- 
oped anteriorly;  last  scale  extending  over  base 
of  caudal  fin.  Gill  rakers  short,  difficult  to  count, 
total  10,  '3  on  upper  arch,  7  on  lower  arch. 
Few  modified  scales  on  snout,  none  on  inter- 
orbit.  Movable  lachrymal  bone  (infraorbital  one) 


FIGURE  3.     Lateral  view  of  holotype  of  Xenaploactis  anopta  (CAS  32633,  37.0  mm  SL). 


POSS  &  ESCHMEYER:  NEW  APLOACTINID  GENUS  AND  SPECIES 


291 


bladelike  with  3  spines:  1st  small,  directed  to- 
ward premaxilla;  followed  by  2  large,  sharp 
spines,  2nd  about  equal  in  length  to  3rd,  di- 
rected downward;  3rd  spine  directed  down 
and  slightly  back.  Spine  on  second  infraor- 
bital  bone  absent;  a  large  elongate  lateral  line 
pore  present.  Mouth  strongly  upturned.  Inter- 
orbit  with  prominent  ridges,  divergent  anterior- 
ly, convergent  over  middle  of  interorbit,  strong- 
ly divergent  posteriorly  (Fig.  4  upper).  Postocular 
spine  and  pterotic  spine  ridgelike.  Posttemporal 
spine  a  large  well-ossified  lump  ending  in  blunt 
spine,  followed  by  blunt  supracleithral  spine. 
Dorsal  posterior  border  of  cleithrum  ending  in 
a  small,  poorly  defined  blunt  spine.  Preopercular 
lateral  line  pores  opening  as  small  tubes,  finger- 
like  cirri  above  uppermost  pores.  In  ventral 
view,  surface  of  dentary  with  5  small  fingerlike 
cirri  along  outer  margin;  5  pairs  of  similar  cirri 
anteriorly,  between  dentaries. 

Color  in  life  unknown.  Color  of  head  and  body 
in  preservative  (Fig.  3)  brown,  with  scattered 
black  specks.  Fins  darker,  possibly  speckled  in 
life;  caudal  with  vertical  bands. 

Measurements  in  millimeters  as  follows  (per- 
cent standard  length  in  parentheses):  standard 
length  37.0;  head  13.0  (35);  snout  3.7  (10);  orbit 
3.3  (9);  interorbital  width  2.5  (6);  jaw  5.3  (14); 
postorbital  7.3  (20);  body  depth  12.3  (33);  pre- 
dorsal  6.4  (17);  anal  fin  14.1  (38);  caudal  fin  9.2 
(25);  pectoral  fin  8.5  (23);  pelvic  fin  5.2  (14);  1st 
dorsal  spine  1.9  (5);  2nd  dorsal  spine  3.5  (9);  3rd 
dorsal  spine  2.4  (6);  4th  dorsal  spine  1.5  (4);  5th 
dorsal  spine  2.2  (6);  penultimate  dorsal  spine  2.2 
(6);  last  dorsal  spine  2.3  (6);  anal  spine  1.9  (5); 
least  depth  of  caudal  peduncle  4.1  (11);  snout  to 
2nd  dorsal  spine  6.8  (18);  snout  to  3rd  dorsal 
spine  8.4  (23);  snout  to  4th  dorsal  spine  12.9  (35); 
snout  to  5th  dorsal  spine  13.7  (37);  width  of  1st 
dorsal  spine  at  midlength  0.3  (1);  incision  of  dor- 
sal fin  membrane  at  4th  dorsal  spine  (from  tip  to 
membrane)  1.5  (4). 

ETYMOLOGY. — The  species-group  name  is  de- 
rived from  the  Greek  anoptos  (unseen). 

DISTRIBUTION. — Known  only  from  the  type- 
locality  in  the  Philippines  at  64-81  m. 

Xenaploactis  cautes,  new  species 

(Figures  4  lower  and  5) 

No  literature  applies  to  this  species. 

MATERIAL.— Hoiotype:  CAS  16105  (28.0  mm  SL).  Gulf  of 
Thailand,  12°19'15"N,  100°43'40"E,  28.6  km  from  Goh 
Chuang,  33  m,  muddy  sand  bottom,  MV  STRANGER,  16-ft  (4.9- 


FIGURE  4.     Dorsal  view  of  head  of  holotypes  of  Xenaploac- 
tis anopta  (upper)  and  X.  cautes  (lower). 


m)  otter  trawl,  George  Vanderbilt  Foundation  sta.  60-449, 
GVF  reg.  no.  2724,  Scripps  locality  60-185  C.N.  633f.5-9a, 
0117-0202  hrs,  13  Dec.  1960.  Paratype:  USNM  221143  (24.3). 
Andaman  Sea,  14°07'N,  97°05'E,  69-73  m,  International  In- 
dian Ocean  Expedition,  ANTON  BRUUN  cruise  1,  sta.  38,  Gulf 
of  Mexico  shrimp  trawl,  30  Mar.  1963. 

COUNTS.— Dorsal  fin  III,  XI,  8-9  (last  dou- 
ble). Anal  fin  I,  10  (last  double).  Pectoral  fin  14. 
Pelvic  fin  I,  3.  Lateral  line  scales  9-10.  Verte- 
brae 27-28. 

DESCRIPTION. — (See  also  generic  description 
above.)  Body  not  notably  elevated  behind  head, 
body  depth  less  than  Vs  of  standard  length.  Body 
covered  with  modified  pointed  scales.  Lateral 
line  with  9-10  tubed  scales,  each  with  2  small 
laterally  projecting  spinules  which  are  best  de- 
veloped anteriorly,  last  scale  extending  over 
base  of  caudal  fin.  Gill  rakers  short,  difficult  to 
count,  total  10-12,  3-4  on  upper  arch,  6-8  on 
lower  arch. 

Many  modified  scales  on  snout.  Lachrymal 
bone  (infraorbital  1)  with  3  spines;  1st,  of  mod- 
erate size,  points  mostly  forward,  continuous 


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10 


FIGURE  5.    Lateral  view  of  holotype  of  Xenaploactis  cautes  (CAS  16105,  28.0  mm  SL). 


with  ridge  at  base  of  larger  2nd  spine;  2nd  spine 
about  equal  in  length  to  3rd,  directed  downward, 
slightly  curved;  3rd  spine  points  mostly  back. 
Second  infraorbital  bone  with  2  spinous  points, 
1  above  other,  with  obscure  circular  lateral  line 
pore.  Interorbit  with  nearly  parallel  ridges, 
stronger  posteriorly  (Fig.  4  lower).  Postocular 
spines  as  slightly  curved  ridges,  meeting  at  mid- 
line  of  interorbit,  connected  to  interorbital 
ridges.  Pterotic  spine  ridgelike.  Posttemporal 
spine  ridgelike,  followed  by  blunt  supracleithral 
spine.  Cleithrum  ending  in  small  blunt  spine.  In 
ventral  view,  surface  of  dentary  with  5  tiny  fin- 
gerlike  cirri  along  outer  margin,  5  pairs  of  similar 
cirri  anteriorly  between  dentaries. 

Color  in  life  unknown.  Color  in  preservative 
pale,  probably  strongly  faded.  Head  and  body 
without  scattered  specks.  Fins  not  darker  than 
body. 

Measurements  in  millimeters  as  follows  (ho- 
lotype first,  percent  standard  length  in  parenthe- 
ses): standard  length  28.0,  24.3;  head  9.9,  9.4 
(35,  39);  snout  2.4,  2.4  (9,  10);  orbit  2.4,  2.4  (9, 
10);  interorbital  width  1.8,  2.1  (6,  9);  jaw  3.6,  4.2 
(13,  17);  postorbit  4.7,  4.9  (17,  20);  body  depth 
7.8,  7.4  (28,  30);  predorsal  5.0,  4.2  (18,  17);  anal 
fin  11.1,  10.8  (40,  44);  caudal  fin  6.7,  6.2  (24,  26); 
pectoral  fin  6.2,  5.5  (22,  23);  pelvic  fin  3.0,  3.4 
(11,  14);  1st  dorsal  spine  1.7,  1.7  (6,  7);  2nd  dor- 
sal spine  2.9,  2.6  (10,  11);  3rd  dorsal  spine  1.7, 
1.6  (6,  6);  4th  dorsal  spine  1.3,  1.3  (4,  5);  5th 
dorsal  spine  1.7,  1.5  (6,  6);  penultimate  dorsal 
spine  2.2,  1.6  (8,  6);  last  dorsal  spine  2.3,  1.4  (8, 


6);  anal  spine  1.8,  2.0  (6,  8);  width  between  in- 
terorbital ridges  0.7,  0.8  (3,  3);  least  depth  of 
caudal  peduncle  3.3,  2.7  (12,  11);  snout  to  2nd 
dorsal  spine  5.6,  5.2  (20,  21);  snout  to  3rd  dorsal 
spine  6.0,  5.4  (21,  22);  snout  to  4th  dorsal  spine 
8.9,  8.4  (32,  34);  snout  to  5th  dorsal  spine  10.3, 
8.5  (37,  35);  width  of  1st  dorsal  spine  at  mid- 
length  0.2,  0.2  (1,  1);  incision  of  fin  membrane 
at  4th  dorsal  spine  (from  tip  to  membrane)  1.3, 
1.3  (4,  5). 

ETYMOLOGY. — The  species-group  name  is  de- 
rived from  the  Latin  cautes  (a  rough,  pointed 
rock)  and  is  to  be  treated  as  a  noun  in  apposi- 
tion. 

DISTRIBUTION. — Known  only  from  the  type 
material  from  the  Andaman  Sea  and  Gulf  of 
Thailand.  This  species  appears  to  inhabit  muddy 
sand  bottom  at  depths  of  33-79  m. 

ACKNOWLEDGMENTS 

We  thank  a  number  of  friends  and  colleagues 
who  assisted  us  in  the  course  of  this  study:  Al- 
wyne  Wheeler  and  Mary  Connolly  (BMNH)  for 
giving  us  the  opportunity  to  examine  the  holo- 
type of  Prosopodasys  asperrimus;  Leslie  Knapp, 
of  the  Smithsonian  Oceanographic  Sorting  Cen- 
ter, for  sending  us  the  second  specimen  of  Xe- 
naploactis cautes;  James  Gordon,  Betty  Powell, 
William  Ruark,  and  Pearl  Sonoda  (all  of  CAS) 
for  their  continous  help.  Tomio  Iwamoto  and 
Lillian  Dempster  (CAS)  reviewed  the  manu- 
script and  offered  critical  suggestions.  We  also 
wish  to  thank  Ellie  Koon  and  Joanne  Zupan,  of 


POSS  &  ESCHMEYER:  NEW  APLOACTINID  GENUS  AND  SPECIES  293 

the  University  of  Michigan,  for  their  assistance.  GUNTHER,  A.  I860.  Catalogue  of  the  acanthopterygian  fishes 
The  drawings  were  made  by  Beth  Meinhard  in  the  British  Museum.  Vol.  2,  Squamipinnes,  Cirrhitidae, 
m  A  c\  Triglidae,  Trachinidae,  Sciaenidae,  Polynemidae,  Trichiur- 

(^t\o).  idae,  Scombridae,  Carangidae,  Xiphiidae.  London,  xxi  + 

548  p. 
LITERATURE  CITED  Poss   §  G    AND  w  N   ESCHMEYER.   1978.  Two  new  Aus- 

CANTOR,  T.   1849.  Catalogue  of  Malayan  fishes.  J.  R.  Asiatic  tralian  velvetfishes,  genus  Paraploactis  (Scorpaeniformes: 

Soc  Bengal  18(2)-  983-1443,  14  pis.  Aploactinidae),  with  a  revision  of  the  genus  and  comments 

ESCHMEYER,  W.  N.   1969.  A  systematic  review  of  the  scor-  on  the  genera  and  species  of  the  Aploactinidae.  Proc.  Calif, 

pionfishes  of  the  Atlantic  Ocean  (Pisces:  Scorpaenidae).  Acad.  Sci.  41(18):401^t26,  14  figs.,  6  tables. 
Occas.  Pap.  Calif.  Acad.  Sci.  79.  130  p. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  9,  pp.  295-302,  6  figs. 


March  5,  1981 


SUNDASALANGIDAE,  A  NEW  FAMILY  OF  MINUTE  FRESHWATER 
SALMONIFORM  FISHES  FROM  SOUTHEAST  ASIA 

By 

Tyson  R.  Roberts 

California  Academy  of  Sciences,  Golden  Gate  Park,  San  Francisco,  CA  94118 


ABSTRACT:  Sundasalangidae,  a  new  family  of  minute  freshwater  salmoniform  fishes  related  to  the  Salangidae 
or  East  Asian  icefishes,  is  based  upon  a  new  genus  and  two  new  species  recently  discovered  in  Southeast  Asia, 
Sundasalanx  praecox  and  S.  microps,  both  described  in  the  present  paper.  With  males  and  females  sexually 
ripe  at  standard  lengths  of  only  14.9  mm,  5.  praecox  is  the  smallest  known  adult  salmoniform  and  is  among  the 
smallest  of  all  adult  vertebrates.  The  new  family  differs  from  all  other  teleosts  including  Salangidae  in  the 
following  features  of  its  skeletal  anatomy,  which  is  largely  cartilaginous:  the  two  halves  of  the  pectoral  girdle 
are  united  to  each  other  by  a  median  scapulocoracoid  cartilage;  in  branchial  arches  1-3  the  basibranchials  and 
hypobranchials  of  either  side  are  represented  by  a  single  cartilaginous  element;  and  each  half  of  the  pelvic 
girdle  is  provided  with  a  pair  of  rod-shaped  parapelvic  cartilages.  Despite  these  and  many  other  differences, 
Sundasalangidae  and  Salangidae  are  clearly  closely  related.  They  agree  with  each  other  but  differ  from  all  other 
known  teleosts  in  having  the  jaw  suspension  with  bilaterally  paired  palatohyomandibuloquadrate  cartilages. 


INTRODUCTION 

Among  the  freshwater  fishes  recently  collect- 
ed by  the  author  in  Southeast  Asia  are  two  sam- 
ples, one  from  far  up  the  mainstream  of  the  Ka- 
puas  River  in  Kalimantan  (Indonesian  Borneo) 
and  the  other  from  a  creek  draining  into  the  Tale 
Sap  in  peninsular  Thailand,  of  minute  transpar- 
ent teleosts  with  a  skeleton  that  is  almost  en- 
tirely cartilaginous  (Figs.  1-2).  Superficially  re- 
sembling drawings  of  early  or  mid-metamorphic 
leptocephalus  larvae  of  Elops  (cf.  Gehringer 
1959:figs.  10-11),  closer  study  of  the  fishes  in 
these  samples  reveals  that  they  are  sexually  ma- 
ture, represent  two  very  distinct  species,  and 
are  actually  salmoniforms  most  closely  related 
to  the  East  Asian  icefishes  of  the  family  Salan- 
gidae. Yet  they  differ  so  markedly  from  Salan- 
gidae, and  in  certain  respects  from  all  other 
known  teleosts,  that  a  new  family  is  proposed 
for  them. 


i        METHODS  AND  MATERIALS; 
ACKNOWLEDGMENTS 

Observations  on  the  largely  cartilaginous  skel- 
etal anatomy  of  Salangidae  and  Sundasalangidae 
have  been  made  on  specimens  prepared  by 
means  of  the  newly  developed  alcian  blue-aliz- 
arin technique  for  counterstaining  cartilage  and 
bone  in  whole,  cleared  specimens  of  small  ver- 
tebrates (Dingerkus  and  Uhler  1977).  I  am  grate- 
ful to  Robert  Drewes  of  the  California  Academy 
of  Sciences  for  preparing  specimens  of  both 
species  of  Sundasalangidae  and  of  all  of  the 
genera  and  nearly  half  of  the  species  of  Salang- 
idae, and  also  to  William  N.  Eschmeyer,  Cu- 
rator of  Fishes  of  the  California  Academy  of 
Sciences,  for  making  the  specimens  of  Salang- 
idae available.  A  complete  list  of  these  skeletal 
preparations  will  be  presented  in  an  exten- 
sively illustrated  account  of  the  skeletal  anat- 
omy of  Salangidae  and  Sundasalangidae  now 


[295] 


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ROBERTS:  SUNDASALANGIDAE— NEW  FISH  FAMILY 


297 


being  prepared  (Roberts,  ms).  I  also  thank  Kar- 
sten  E.  Hartel  and  William  Fink  for  loaning  the 
specimens  of  Sundasalanx  praecox  which  had 
been  deposited  by  me  in  the  Museum  of  Com- 
parative Zoology  at  Harvard.  My  fieldwork  in 
Thailand  and  Indonesia  which  led  to  the  discov- 
ery of  Sundasalanx  was  supported  or  aided  by 
the  following  institutions:  Museum  of  Compar- 
ative Zoology;  Kasetsart  University  College  of 
Fisheries;  National  Research  Council,  Thailand; 
Indonesian  National  Research  Council  (LIPI); 
and  Smithsonian  Tropical  Research  Institute 
(STRI).  I  particularly  thank  Supap  Monkolprasit 
of  Kasetsart  and  Ira  Rubinoff  of  STRI  for  facil- 
itating my  travel  and  fieldwork.  The  preparation 
of  this  paper  was  supported  by  grant  DEB77- 
24574  in  the  Systematic  Biology  Program  of  the 
National  Science  Foundation. 

SUNDASALANGIDAE,  new  family 

TYPE-GENUS:  Sundasalanx,  new  genus. 

Sundasalangids  differ  from  all  other  teleosts, 
so  far  as  known,  in  having  a  pectoral  girdle  with 
a  median  cartilaginous  scapulocoracoid  and  a 
single  pair  (one  on  each  side)  of  fan-shaped, 
"externalized"  radial  cartilages  (Fig.  3)  which 
form  the  peduncular  portion  of  the  pedunculated 
pectoral  fins;  a  pair  of  rod-shaped  parapelvic 
cartilages  (Fig.  2)  which  apparently  serve  to 
anchor  each  half  of  the  pelvic  girdle  to  the  free 
ventral  end  of  a  myotomal  muscle;  and  branchial 
arches  with  hypobranchial  elements  1-3  absent 
as  separate  elements,  evidently  fused  to  basi- 
branchials  1-3  (Fig.  4).  In  Salangidae  the  scap- 
ulocoracoids  are  paired,  separate  elements,  and 
the  two  halves  of  the  pectoral  girdle  are  separate 
from  each  other;  each  pectoral  fin  is  supported 
by  three  or  more  radial  cartilages;  parapelvic 
cartilages  are  absent;  and  the  first  three  bran- 
chial arches  have  separate  basibranchial  and  hy- 
pobranchial elements. 

Members  of  the  Sundasalangidae  agree  with 
most  Salangidae  but  apparently  differ  from  all 
other  teleosts  in  having  a  jaw  suspension  con- 
sisting of  a  single  cartilaginous  element  or  pal- 
atohyomandibuloquadrate  (Fig.  5);  they  agree 
with  Salangidae  but  apparently  differ  from 
adults  of  all  other  known  teleosts  (Nelson 
1960:61)  in  having  well-developed  separate 
fourth  hypobranchials  (Fig.  4).  Sundasalangids 
agree  with  the  Salangidae  but  differ  from  adults 
of  most  other  teleosts  in  having  pedunculate 
pectoral  fins;  a  scaleless  body;  no  symplectics; 


no  circumorbital  bones;  myotomal  muscles  fail- 
ing to  meet  at  ventral  midline  of  body;  and  max- 
illary bones  with  distal  two-thirds  curved  in- 
wards underneath  the  head  so  that  the  portion 
of  the  maxillary  toothrow  they  bear  projects 
medially  rather  than  ventrally  when  the  mouth 
is  closed.  In  addition  to  the  unique  characters 
associated  with  their  pectoral  and  pelvic  girdles 
and  gill  arches,  which  have  been  described 
above,  Sundasalangidae  differ  from  Salangidae 
in  their  much  smaller  size  (the  smallest  salangids 
are  over  35  mm  in  standard  length  [SL]  when 
sexually  mature);  olfactory  organs  each  with  a 
single  nasal  opening  instead  of  two  openings; 
interopercle  absent;  pectoral  fin  without  seg- 
mented bony  rays;  adipose  fin  absent;  sexually 
mature  males  without  enlarged  or  otherwise 
modifed  anal  fin,  or  a  row  of  large  pored  scales 
on  the  base  of  the  anal  fin  (present  in  all  salan- 
gids); pelvic  fins  five-rayed  (seven-rayed  in  all 
salangids);  myotomes  <-shaped  (£ -shaped  in 
salangids);  and  vertebrae  only  37-43  (48-77  in 
Salangidae).  Sundasalangidae  comprises  two 
species,  Sundasalanx  praecox  and  S.  microps, 
described  below. 

Sundasalanx,  new  genus 

TYPE-SPECIES:  Sundasalanx  praecox,  new  species. 

Minute,  transparent,  freshwater  fishes  with  a 
largely  cartilaginous  skeleton;  a  single  large  na- 
sal opening  on  each  side  of  snout;  body  bilat- 
erally compressed,  scaleless;  branchiostegal 
rays  four  (usually  three  in  Salangidae);  pectoral 
fins  pedunculate,  without  bony  rays;  pelvic  fins 
midabdominal,  with  five  rays;  myotomal  mus- 
culature of  opposite  sides  widely  separated  ven- 
trally; each  half  of  pelvic  girdle  anchored  to  free 
ventral  end  of  a  myotomal  muscle  by  a  pair  of 
parapelvic  cartilages;  a  median  membranous 
keel  extending  on  abdomen  from  pelvic  fins  to 
vent;  gut  a  simple  straight  tube  with  no  differ- 
entiated stomach;  vent  immediately  anterior  to 
anal  fin  origin;  no  secondary  sexual  dimorphism 
or  dichromatism;  vertebrae  (including  hypural 
centrum  as  one)  37-43. 

Dentition  (Figs.  4-5):  Premaxillary  and  max- 
illary with  a  single  row  of  minute  conical  teeth; 
lower  jaw  with  two  rows  of  minute  conical  teeth, 
inner  row  curved  inwards  away  from  outer  row; 
fifth  ceratobranchials  with  0-10  conical  teeth. 
Roof  of  mouth  and  upper  pharyngeal  elements 
edentulous. 

Median  fins:  Dorsal  and  anal  fins  originating 


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299 


on  posterior  third  of  body;  origin  of  anal  fin  on 
a  vertical  through  or  slightly  posterior  to  middle 
of  dorsal  fin  base.  Dorsal  fin  with  three  simple 
and  8-11  branched  rays  (last  ray,  counted  as 
one,  split  to  base);  anal  fin  with  3-4  simple  rays 
and  12-17  branched  rays  (last  ray  split  to  base). 
Adipose  fin  absent.  Caudal  fin  moderately 
forked,  with  10  +  9  principal  rays  and  9-12  + 
7-11  procurrent  rays. 

ETYMOLOGY. — Sunda  (=Sundaland,  the  con- 
tinental landmass  of  Southeast  Asia  connected 
to  the  Asian  mainland  by  the  isthmus  of  Kra) 
plus  Salanx  (Greek,  masc.),  type-genus  of  the 
family  Salangidae. 

Sundasalanx  praecox,  new  species 

(Figures  la,  2a,  4a,  5a,  6a) 

MATERIAL.— Holotype:  MCZ  47129,  17.2-mm  male  with 
well-developed  testes,  from  Khlong  Falamee,  a  swift,  muddy 
creek  1-2  m  deep  and  3-5  m  wide  with  hard-packed  mud 
bottom  flowing  into  inner  lake  of  Tale  Sap,  at  about  2  km  W 
of  Pak  Payoon  on  the  isthmus  of  Kra,  southern  Thailand;  ny- 
lon flyscreen  pushnet;  20  June  1970. 

Paratypes:  MCZ  54390,  1 19:  14.9-18.3  mm,  same  collection 
data  as  holotype;  nine  utilized  for  alcian  blue-alizarin  prepa- 
rations. 

DIAGNOSIS. — S.  praecox  is  distinguished  from 
S.  microps,  its  only  congener,  by  its  much  larger 
eyes:  horizontal  diameter  of  eye  measured  in  10 
specimens  including  smallest  and  largest  speci- 
men of  each  species,  4.3-5.0%  of  SL  in  S.  prae- 
cox vs.  2.7-3.2%  in  S.  microps.  In  S.  praecox 
eyeballs  separated  from  each  other  by  a  distance 
about  equal  to  transverse  diameter  of  their  pig- 
mented  portion,  while  in  5.  microps  distance 
separating  them  equal  to  at  least  twice  trans- 
verse diameter.  In  S.  praecox  head  deeper, 
more  compressed,  and  nasal  septum  much  nar- 
rower (Figs.  1,  6).  Maxillary  teeth  about  15-19 
vs.  about  30  (Fig.  5).  Palatohyomandibuloquad- 
rate  cartilage  entire  vs.  palatine  separate  from 
hyomandibuloquadrate  (Fig.  5).  Fifth  cerato- 
branchial  with  about  8-10  large  conical  teeth  vs. 
0-3  small  conical  teeth  (Fig.  4).  First  gill  arch 
with  1  +  9  well-developed  gill  rakers  vs.  0+1- 
2  rudimentary  gill  rakers;  all  ceratobranchials 
with  well-developed  gill  rakers  vs.  gill  rakers 
greatly  reduced  in  size,  and  ceratobranchials 
moderately  elongate  vs.  slender  and  very  elon- 
gate (Fig.  4).  Posterior  parapelvic  cartilage  orig- 
inating dorsal  to  anterior  parapelvic  cartilage 
and  extending  further  into  myotomal  muscle 
mass  instead  of  lying  parallel  with  anterior  par- 
apelvic cartilage  (Fig.  2).  No  midventral  row  of 


large,  round  pigment  spots  on  abdomen  and 
postpelvic  abdominal  keel  corresponding  in 
number  to  myotomal  muscles  (present  in  5.  mi- 
crops)  (Fig.  1).  Anal  fin  with  12-15  branched 
rays  vs.  14-17.  Caudal  peduncle  more  slender 
(Figs.  1,  2).  Vertebral  centra  more  elongate  (Fig. 
2.)  Total  vertebrae  (excluding  basioccipital  half 
centrum  but  counting  upturned  hypural  centrum 
as  one)  37(n  =  2)  or  38(7)  vs.  41(2),  42(4)  or 
43(1). 

Sex:  The  120  5.  praecox  from  Khlong  Fala- 
mee were  all  caught  in  a  segment  of  the  creek 
less  than  100  m  long  and  presumably  represent 
a  random  sample  from  a  single  breeding  popu- 
lation. The  sample  includes  32  males  14.9-18.3 
mm  SL  with  well-developed  testes,  19  females 
14.9-17.3  mm  with  well-developed  eggs,  and  68 
specimens  15.2-17.9  mm  of  undetermined  sex 
in  which  the  gonads  are  inactive  or  relatively 
undeveloped.  The  testes  extend  nearly  the  entire 
length  of  the  abdomen  dorsolaterally  to  the  gut. 
Obscured  anteriorly  by  the  liver,  the  testes  are 
otherwise  readily  visible  through  the  transparent 
ventral  body  wall  and  translucent  ventral  por- 
tion of  the  myotomal  muscles  (Fig.  la);  they  are 
uniformly  divided  throughout  their  length  into 
obliquely  aligned  divisions  or  partitions,  of 
which  there  are  about  five  or  six  per  myotome. 
Similarly  partitioned  testes,  present  in  some 
minute  or  small  freshwater  African  Clupeidae 
(personal  observation  with  Peter  Whitehead) 
have  not  been  observed  in  Salangidae.  The  larg- 
est specimen  in  the  entire  sample  is  a  male  of 
18.3  mm  with  very  well  developed  testes;  the 
smallest  male,  14.9  mm,  has  testes  almost  as 
well  developed.  In  females  the  ovaries  also  ex- 
tend virtually  the  entire  length  of  the  abdomen 
and  lie  dorsolateral  to  the  gut.  Eggs  in  varying 
stages  of  development  are  present.  In  the  ripest 
ovaries  observed,  the  largest  eggs,  0.20-0.25 
mm  in  diameter,  are  aligned  in  a  single  row  of 
about  25  eggs  in  each  ovary.  The  smallest  fe- 
male, 14.9  mm,  has  ovaries  with  well-developed 
eggs. 

Food:  Gut  contents,  either  whole  macro- 
scopic animals  or  fragments  of  them,  readily  ob- 
served through  the  transparent  body  and  gut 
walls,  are  present  in  34  (28%)  of  the  120  speci- 
mens. In  the  other  86  (72%),  the  guts  appear  to 
be  entirely  empty.  Items  ingested  consist  exclu- 
sively of  animals,  almost  all  apparently  either 
aquatic  insect  larvae  or  segmented  vermiform 
organisms  (also  aquatic  insects?).  None  of  the 


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PT 


FIGURE  3.  Sundasalanx  microps,  CAS  44220,  17.0  mm, 
pectoral  girdle  and  fin  (ventral  view).  PT,  SCL,  CL  =  post- 
temporal,  supracleithrum,  cleithrum;  R,  F  =  pectoral  radial, 
frayed  margin  of  fin;  SCO  =  scapulocoracoid  (secondary  pec- 
toral girdle  and  frayed  margin  of  fin  of  right  side  omitted). 


guts  contain  plant  material,  sediment,  or  signif- 
icant amounts  of  nonidentifiable  debris. 

ETYMOLOGY. — Latin  praecox,  too  early  ripe, 
premature. 


Sundasalanx  microps,  new  species 

(Figures  Ib,  2b,  3,  4b,  5b,  6b) 

MATERIAL. — Holotype:  Museum  of  Zoology,  Bogor,  Indo- 
nesia, 3000,  17.0  mm,  near  shore  of  mainstream  Kapuas 
River  at  Kampong  Nibung,  about  100  km  NE  of  Sintang  and 
7  km  NE  of  Selimbau,  Kalimantan,  Indonesia,  lat.  0°39'N, 
long.  112°10.5'E;  current  moderate,  water  muddy,  26°C,  pH 
5.5-6,  bottom  soft  mud,  depth  to  1  m;  nylon  flyscreen  seine; 
5-6  July  1976. 

Paratypes:  Museum  of  Zoology,  Bogor,  Indonesia,  3001, 
and  CAS  44220,  34:14.6-19.9  mm,  same  collection  data  as 
holotype,  seven  utilized  for  alcian  blue-alizarin  preparations. 

DIAGNOSIS. — S.  microps  is  distinguished  from 
S.  praecox,  its  only  known  congener,  in  the  di- 
agnosis of  that  species  given  above. 

Sex  and  food:  The  gonads  in  the  sample  of  35 
S.  microps  are  not  well  developed,  and  I  have 
been  unable  to  distinguish  males  and  females. 
The  guts  of  nearly  all  appear  to  be  empty;  a  few 
contain  unidentified  fragments,  but  no  whole  in- 
sect larvae  or  other  animals,  plant  material,  or 
sediment. 

ETYMOLOGY. — Greek  mikros,  small,  little, 
and  ops,  eye. 

Sundasalanx  species  undetermined 

Vaillant  (1893:110-112,  pi.  2,  fig.  4)  described 
some  delicate  little  fishes  collected  in  the  Ka- 


1  mm 


B,  H  1-2 


C1-5 


FIGURE  4.  Ventral  parts  of  gill  arches,  dorsal  view  (gill  rakers  of  right  side  omitted):  (a)  Sundasalanx  praecox,  MCZ  54390, 
17.2  mm;  (b)  Sundasalanx  microps,  CAS  44220,  17.2  mm.  B,  H,  C  =  basibranchial,  hypobranchial,  ceratobranchial  (see  text 
for  explanation  ). 


ROBERTS:  SUNDASALANGIDAE— NEW  FISH  FAMILY 


301 


OP 


LJ 


MX 


PL        HQ 


FIGURE  5.  Jaws,  jaw  suspension,  and  gill  cover,  lateral  view:  (a)  Sundasalanx  praecox,  MCZ  54390,  17.1  mm;  (b)  Sun- 
dasalanx  microps,  MCZ  44220,  17.0  mm.  P,  MX,  LJ  =  premaxillary,  maxillary,  lower  jaw;  PHQ  =  palatohyomandibuloquad- 
rate;  PL,  HQ  =  palatine,  hyomandibuloquadrate;  OP,  SO  =  opercle,  subopercle. 


puas  River  by  Chaper  in  1890-91  which  he  hes- 
itantly identified  as  young  needlefish  (Welone 
caudimaculatd).  Subsequent  to  completing  the 
manuscript  of  this  paper,  I  visited  the  Museum 
National  d'Histoire  Naturelle  in  Paris,  where 
Chaper' s  Kapuas  collection  is  deposited,  and 
examined  this  material.  The  specimens  (MNHN 
91-596,  27:18.8-23.4  mm),  although  not  in  the 
best  state  of  preservation,  are  clearly  Sundasa- 
lanx but  do  not  agree  well  with  my  diagnoses  of 
either  5.  microps  or  5.  praecox.  They  agree 
with  5.  microps  rather  than  S.  praecox  in  their 
relatively  large  size,  vertebral  counts  of  about 
42-45,  modally  43  (determined  by  counting  the 
myotomes),  and  number  of  branched  anal  fin 
rays,  about  15-18.  On  the  other  hand,  the  eyes 
seem  to  be  much  larger  than  in  5.  microps  and 
possibly  even  larger  than  in  5.  praecox,  and  the 
number  of  maxillary  teeth  fewer  than  in  S.  mi- 
crops  and  5.  praecox.  In  nearly  all  of  the  spec- 
imens, the  eyes  are  ruptured  or  their  shape  so 
distorted  that  horizontal  diameter  cannot  be 
measured  accurately.  In  a  22.5-mm  specimen  in 
which  the  eye  is  intact  and  nearly  normal  in 
shape,  its  horizontal  diameter  is  5.0%  of  SL 
(2.7-3.2%  in  S.  microps,  4.3-5.0%  in  5.  prae- 
cox). The  eyeballs  are  so  large  that  they  nearly 
meet  in  the  middle  of  the  head,  whereas  in  both 
S.  microps  and  S.  praecox  they  are  consider- 


ably further  apart.  The  number  of  teeth  on  the 
maxillary  bone  is  only  10-14  (n  =  3)  vs.  about 
30  in  S.  microps  and  about  15-19  in  5.  praecox. 
A  row  of  melanophores  is  visible  on  the  side  of 
the  body  external  to  the  free  ventral  ends  of  the 
myotomal  muscles,  but  not  on  the  ventral  mid- 
line  of  the  abdomen  and  postpelvic  abdominal 
keel  (the  latter  present  in  S.  microps  but  not  in 
S.  praecox).  I  was  unable  to  detect  testes  or 
eggs  in  any  of  the  specimens,  although  many  of 


ACF 


FIGURE  6.  Anterior  end  of  cranium,  dorsal  view:  (a)  Sun- 
dasalanx praecox,  MCZ  54390,  17.2  mm;  (b)  Sundasalanx 
microps,  CAS  44220,  17.0  mm.  EP,  NS  =  ethmoid  plate,  na- 
sal septum;  ACF,  TC  =  anterior  cranial  fontanel,  tectum  cra- 
nii. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  9 


them  have  ruptured  abdomens.  The  precise  lo- 
cality where  they  were  collected  is  unknown, 
but  judging  from  Vaillant's  remarks  (1893:60) 
they  must  have  been  taken  in  or  near  the  main- 
stream of  the  Kapuas  River  somewhere  between 
its  confluence  with  the  Sebruang  River  and  Sem- 
itau,  or  in  the  same  general  area  as  S.  microps, 
the  type-locality  of  which  is  less  than  40  km 
upriver  from  Semitau.  So  far  as  I  have  been  able 
to  find  out,  no  additional  material  of  Sundasa- 
lanx  has  been  reported  upon  or  collected  by  any- 
one else. 

DISCUSSION 

Alizarin  is  a  specific  stain  for  bone  but  does 
not  always  stain  bone  in  an  early  stage  of  de- 
velopment or  poorly  calcified  bone.  Alcian  blue 
usually  stains  cartilage,  but  apparently  also 
sometimes  stains  uncalcified  bone  or  bone  in  an 
early  stage  of  development.  In  some  instances 
structures  which  are  clearly  bone  or  cartilage 
fail  to  be  noticeably  stained  by  either  alizarin  or 
alcian  blue.  Thus,  it  is  not  possible  to  state  in 
every  instance  which  skeletal  elements  in  Sun- 
dasalangidae  and  Salangidae  are  cartilage  and 
which  are  bone  based  merely  on  their  staining 
reaction  to  alcian  blue  and  alizarin.  In  Sunda- 
salangidae  the  only  skeletal  elements  deeply 
stained  with  alizarin  are  vertebral  centra,  distal 
ends  of  neural  and  haemal  spines,  caudal  fin 
rays,  hypural  fan,  teeth,  and  bony  toothplates 
on  fifth  ceratobranchials.  Maxillary  bone,  tooth- 
bearing  portion  of  the  lower  jaw,  secondary  pec- 
toral fin  girdle  (posttemporal,  supracleithrum 
and  cleithrum)  and  pelvic  fin  rays  are  weakly 
stained  with  alizarin.  Gill  rakers,  branchiostegal 
rays,  and  subopercle  are  either  weakly  stained 
with  alcian  blue  or  are  not  stained  at  all,  in  which 
case  they  may  be  difficult  to  observe  even  with 
transmitted  light.  In  alcian  blue-alizarin  prepa- 
rations of  adult  Salangidae,  the  distribution  of 


elements  stained  by  alcian  blue  and  alizarin  is 
basically  similar  to  that  in  Sundasalangidae,  but 
alizarin  is  taken  up  more  extensively. 

Sundasalanx  praecox  is  among  the  smallest  of 
adult  vertebrates  and  is  the  smallest  known  sal- 
moniform  fish.  Sundasalanx  microps  is  the 
smallest  of  more  than  250  fish  species  present  in 
the  Kapuas  River  (personal  observations),  thus 
providing  an  excellent  example  of  survival  of  a 
peripheral-division  fish  family  in  the  midst  of  a 
rich  primary-division  freshwater  ichthyofauna 
by  evolution  of  minute  body  size  and  a  wholly 
freshwater  life  history  (for  further  discussion  see 
Roberts  1978:20-21).  Salangidae  are  anadro- 
mous  and  freshwater  fishes  inhabiting  coastal 
waters  and  rivers  of  East  Asia  from  North  Viet- 
nam northwards  to  Korea,  Vladivostok,  and 
Sakhalin.  They  are  unknown  from  Thailand  and 
Borneo.  The  basic  references  to  systematics  and 
geographical  distribution  of  Salangidae  are  Fang 
(1934)  and  Wakiya  and  Takahasi  (1937). 

LITERATURE  CITED 

DINGERKUS,  G.,  AND  L.  D.  UHLER.  1977.  Enzyme  clearing 
of  alcian  blue  stained  whole  small  vertebrates  for  demon- 
stration of  cartilage.  Stain  Tech.  52(4): 229-232,  3  figs. 

FANG,  P.  W.  1934.  Study  on  the  fishes  referring  to  Salangidae 
of  China.  Sinensis  4(9):231-268,  7  figs.,  6  tables. 

GEHRINGER,  J.  W.  1959.  Early  development  and  metamor- 
phosis of  the  ten-pounder,  Elops  saurus  Linnaeus.  U.S. 
Fish  Wildl.  Serv.  Fish.  Bull.  59  (155):619-647,  32  figs.,  14 
tables. 

NELSON,  G.  J.  1970.  Gill  arches  of  some  teleostean  fishes  of 
the  families  Salangidae  and  Argentinidae.  Jpn.  J.  Ichthyol. 
17(2):61-66,  2  figs. 

ROBERTS,  T.  R.  1978.  An  ichthyological  survey  of  the  Fly 
River  in  Papua  New  Guinea  with  descriptions  of  new 
species.  Smithson.  Contrib.  Zool.  281,  72  pp.,  32  figs. 

VAILLANT,  L.  1893.  Contribution  a  I'etude  de  la  faune  ichth- 
yologique  de  Borneo.  Nouv.  Arch.  Mus.  Hist.  Nat.,  ser.  3, 
5:23-114,  2  pis. 

WAKIYA,  Y.,  AND  N.  TAKAHASI.  1937.  Study  on  fishes  of 
the  family  Salangidae.  J.  Coll.  Agric.  Tokyo  Univ. 
14(4): 265-295,  3  figs.,  pis.  16-21,  2  tables. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  10,  pp.  303-314,  4  figs. 


March  5,  1981 


NEW  AND  RECONSIDERED  SPECIES  OF  MICONIA 

(MELASTOMATACEAE)  FROM 

COSTA  RICA  AND  PANAMA 

By 
Frank  Almeda 

Department  of  Botany,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  94118 


ABSTRACT:  Diagnoses,  descriptions,  and  discussions  are  presented  for  five  new  Central  American  species  of 
Miconia  (M.  chiriquiensis  and  M .  coloradensis  from  Panama,  and  M.  confertiflora,  M.  grandidentata,  and  M. 
longibracteata  from  Costa  Rica).  Diagnostic  illustrations  are  provided  for  four  of  the  novelties,  and  the  new 
name  Miconia  concinna  is  proposed  for  a  Panamanian  endemic  heretofore  known  as  Topobea  micrantha. 


INTRODUCTION 

In  the  four  decades  since  Standley  (1938)  at- 
tempted to  summarize  knowledge  of  the  Costa 
Rican  Melastomataceae,  increased  botanical  ex- 
ploration in  Costa  Rica  and  adjacent  Panama  has 
resulted  in  the  collection  of  much  new  material 
of  the  montane  species  of  Miconia.  Field  and 
herbarium  study  of  this  genus  for  a  comprehen- 
sive treatment  of  Melastomataceae  for  Flora 
Costaricensis  reveals  the  need  for  nomenclatur- 
al  changes,  emended  species  descriptions,  and 
recognition  of  several  undescribed  taxa.  This 
paper  places  some  of  this  new  information  on 
record  now  to  make  the  names  available  to  other 
researchers  prior  to  completion  of  the  floristic 
treatment. 

Miconia  chiriquiensis  Almeda,  sp.  nov. 

(Figure  1) 

Sect.  Cremanium.  Frutex  vel  arbor  parva  2- 
4(-IO)  m.  Ramuli  teretes  vel  obscure  rotundato- 
quadrangulati  sicut  folia  primum  paulo  furfu- 
racei  mox  glabrati.  Petioli  (5-)7-18(-25)  mm; 
lamina  3. 7-7. 5( -10)  x  1.5-4.6  cm  chartacea  el- 


liptica  apice  caudato-acuminato  basi  acuta 
margine  obscure  serrulato.  Panicula  4-6.5  cm 
longa  glabra  laxe  multiflora;  flores  5-meri,  pedi- 
cellis  0.5-1  mm  longis,  bracteolis  0.5-1 .5  x  0.5 
mm  caducis.  Hypanthium  (ad  torum)  I  x  I  mm, 
lobis  interioribus  0.5-1  mm  altis  late  deltoideis 
apice  obtuso,  dentibus  exterioribus  acutis  ca. 
0.5  mm  eminentibus.  Petala  1.5  x  1.5  mm  sub- 
orbicularia  glabra.  Stamina  isomorphica  gla- 
bra; filamenta  1 .5  mm  longa;  antherarum  thecae 
0.75  x  0.5  mm  paulo  cuneatae  poro  0.5  mm 
diam.  ventraliter  inclinato,  connectivo  non  pro- 
longato.  Stylus  0.7-1  x  0.5  mm  glaber;  stigma 
subcapitatum;  ovarium  omnino  inferum  apice 
glabro. 

Shrub  or  tree  2^(-10)  m  tall.  Internodes  and 
distal  branches  ±  terete,  essentially  glabrous  at 
maturity,  but  vegetative  buds  and  young  leaves 
commonly  beset  with  a  brownish  furfuraceous 
indument.  Leaves  chartaceous,  distantly  ciliate- 
serrulate,  3.7-7.5(-10)  cm  long  and  1.5-4.6  cm 
wide,  elliptic,  bluntly  caudate-acuminate  apical- 
ly  and  acute  basally,  glabrous  at  maturity, 


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ALMEDA:  NEW  SPECIES  OF  MICONIA 


305 


3-nerved  or  3-plinerved  (excluding  inconspic- 
uous submarginal  pair)  punctate  and  with  a  con- 
spicuous network  of  secondaries  below;  petioles 
(5_7)7_18(-25)  mm  long  and  1  mm  broad.  Inflo- 
rescence a  laxly  branched  suberect  to  ±  pendant 
panicle  4-6.5  cm  long;  rachis  ±  quadrate,  gla- 
brous throughout;  bracteoles  sessile,  early-de- 
ciduous, 0.5-1.5  mm  long  and  0.5  mm  wide,  lin- 
ear-oblong, entire.  Pedicels  0.5-1  mm  long. 
Hypanthia  (at  anthesis)  campanulate,  1  mm  long 
to  the  torus,  glabrous.  Calyx  lobes  (on  fruiting 
hypanthia)  persistent,  glabrous,  semicircular  or 
depressed-triangular  with  entire  to  irregularly, 
±  hyaline  margins,  0.5-1  mm  long  and  about  1 
mm  wide;  calyx  teeth  persistent,  subulate,  ± 
appressed  to  and  shorter  than  mature  calyx 
lobes.  Petals  5,  erect  to  antrorsely  spreading, 
glabrous,  ±  concave,  white  or  greenish  white, 
suborbicular,  entire,  1 .5  mm  long  and  wide.  Sta- 
mens 10,  isomorphic,  incurved  toward  central 
axis  of  flower  at  anthesis;  filaments  glabrous, 
subulate,  white  to  translucent,  mostly  1.5  mm 
long  and  0.5  mm  wide  basally;  anthers  about 
0.75  mm  long  and  0.5  mm  wide  distally,  white, 
±  infundibuliform  to  obliquely  cuneate  in  profile 
view,  shallowly  emarginate  dorsally,  the  pore 
oblong  and  ventrally  inclined;  connective  sim- 
ple, inconspicuous  and  lacking  prolongations  or 
appendages.  Ovary  inferior.  Style  straight,  gla- 
brous, 0.7-1  mm  long;  stigma  subcapitate.  Berry 
globose,  2-3  mm  long  to  the  torus  and  2-3.5  mm 
in  diameter.  Seeds  ±  pyriform,  beige,  papillate, 
mostly  0.7-1  mm  long. 

TYPES. — Panama.  Chiriqui:  about  8  km  w  of  Cerro  Punta, 
vicinity  of  Las  Nubes,  elevation  6100-6400  ft  [1859-1951  m], 
1 1  Feb.  1978,  Almeda  &  Nakai  3535  (holotype:  CAS!;  iso- 
types:  C!,  DUKE!,  F!,  MO!,  PMA!,  US!). 

ADDITIONAL  SPECIMENS  EXAMINED. — Panama.  Chiriqui: 
vicinity  of  Las  Nubes,  2.7  miles  [4.3  km]  NW  of  Rio  Chiriqui 
Viejo,  w  of  Cerro  Punta,  Croat  22392  (CAS,  DUKE,  MO), 
Croat  22425  (MO,  US);  Bajo  Chorro,  Boquete,  Davidson  181 
(US),  Davidson  390  (MO). 

DISTRIBUTION. — A  little-collected  cloud-for- 
est species  apparently  restricted  to  Chiriqui 
province  in  western  Panama  at  elevations  of 
1850-2200  m.  Available  specimens,  all  of  which 


were  collected  in  February,  are  in  flower  and 
fruit. 

Miconia  chiriquiensis  is  apparently  rare  and 
occurs  in  a  region  which  continues  to  yield  new 
and  narrowly  endemic  taxa.  Diagnostic  features 
include  the  ciliate-serrulate,  elliptic  leaves  that 
are  caudate-acuminate  apically,  short  (0.5-1 
mm),  linear-oblong,  early-deciduous  bracteoles, 
concave,  suborbicular  petals,  geniculate  fila- 
ments, and  minute  (0.75  x  0.5  mm),  4-celled  an- 
thers. 

In  his  account  of  Miconia  for  the  Melasto- 
mataceae  of  Panama,  Gleason  (1958)  referred 
this  entity  to  M.  rubens  (Sw.)  Naud.  Study  of 
his  description  and  examination  of  selected  cited 
specimens  indicate  that  Gleason  also  confused 
M.  chiriquiensis  with  the  taxon  treated  here  as 
M.  concinna.  The  latter  differs  markedly  from 
M.  chiriquiensis  by  virtue  of  its  epiphytic  habit, 
trichotomously  branched,  corymbiform  panicle, 
and  distinctive  androecial  morphology.  In  foliar 
size  and  shape  the  new  species  bears  a  strong 
resemblance  to  M.  rubens,  which  is  known  only 
from  Jamaica  and  Venezuela.  Miconia  rubens 
does  differ  conspicuously,  however,  by  the 
somewhat  swollen  nodes,  ferrugineous  pubes- 
cence on  distal  nodes  and  juvenile  foliage,  mar- 
ginally fimbriate  bracteoles,  dioecious  floral 
condition,  peltate  stigma,  and  cuneate,  apically 
truncate  anthers.  Although  M.  chiriquiensis  re- 
sembles M.  rubens  most  closely  in  the  totality 
of  its  vegetative  characters,  it  is  difficult  to  pres- 
ent meaningful  speculation  regarding  the  origin 
and  exact  relationships  of  these  taxa.  Aside  from 
the  possibility  of  evolutionary  convergence,  the 
most  logical  alternative  hypothesis  is  that  M. 
rubens  is  a  close  relative  and  possibly  dioecious 
derivative  of  M.  chiriquiensis. 

Miconia  coloradensis  Almeda,  sp.  nov. 

(Figure  2) 

Sect.  Amblyarrhena.  Herba  I  m  alt  a  (fide  col- 
lectore).  Ramuli  glabri  primum  obscure 
subquadrangulati  demum  terete s.  Petioli  3- 
7.5  x  i.5-3  cm;  lamina  13.5-20.5  x  9.8-17  cm 


FIGURE  1.  Miconia  chiriquiensis  Almeda.  A,  habit,  x'/i;  B,  representative  leaves,  lower  surface  (left)  and  upper  surface 
(right),  xl;  C,  seeds,  x!2;  D,  mature  berry,  x8;  E,  stamens,  ventral  view  (left)  and  lateral  view  (right)  x  ca.  10;  F,  petal, 
x  ca.  8;  G,  fully  expanded  flower  showing  natural  posture  of  petals  and  stamens  (left),  floral  bud  and  pedicellar  bracteoles  (right), 
x9.  (A-G  from  Almeda  &  Nakai  3535.) 


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chartacea  vel  subcoriacea  denticulata  cordata 
apice  acuminato  vel  cuspidato,  9-1 1-nervata, 
supra  glabra  et  paulo  reticulato-bullata,  subtus 
in  superficie  pilis  stellatis  obsita.  Panicula  7-19 
cm  longa  multiflora;flores  (4-)5-meri,  1 .5-5  mm 
pedicellati,  bracteolis  3-6  mm  longis  obovatis 
vel  spathulatis  valde  caducis.  Hypanthium  (ad 
torum)  2.5  x  2  mm,  lobis  interioribus  I  mm  altis 
rotundatis  vel  truncatis,  dentibus  exterioribus 
crassis  ca.  0.5  mm  eminentibus.  Petala  3-5.5  x 
2-4  mm  obovata.  Stamina  isomorphica  glabra; 
filamenta  2.5  mm  longa;  antherarum  thecae 
2.5  x  1  mm  anguste  obovatae  poro  0.5-0.75 
mm  diam.  paulo  ventraliter  inclinato;  connec- 
tivum  nee  prolongatum  nee  appendiculatum. 
Stylus  5  x  0.5  mm  glaber;  stigma  subcapita- 
tum. 

Herb  to  1  m  tall  (according  to  collectors). 
Cauline  internodes  glabrous,  subquadrangular  to 
±  terete.  Leaves  ±  bullate  above,  chartaceous 
to  subcoriaceous,  denticulate,  13.5-20.5  cm  long 
and  9.8-17  cm  wide,  cordate,  acuminate  to  cus- 
pidate apically,  9-1 1 -nerved  with  a  prominulous 
network  of  secondary  and  tertiary  nerves  below, 
glabrous  and  green  above,  red  to  purple  and 
moderately  beset  with  sessile  stellate  trichomes 
on  and  between  primary  and  secondary  nerves 
below;  petioles  3-7.5  cm  long  and  1.5-3  cm 
broad,  each  petiole  bearing  an  abaxial,  humplike 
protuberance  proximal  to  nodal  junction.  Inflo- 
rescence a  laxly  branched  terminal  panicle  7-19 
cm  long;  rachis  ±  rounded  to  subquadrangular, 
moderately  stellate  pubescent;  bracteoles  ses- 
sile, early  deciduous,  3-6  mm  long  and  1.5-3 
mm  wide,  obovate  to  spatulate,  erose  to  dentic- 
ulate, ±  enveloping  young  buds  and  pedicels, 
glabrous  above,  sparsely  to  moderately  stellate 
below.  Pedicels  1.5-5  mm  long,  beset  with  ses- 
sile stellate  trichomes.  Hypanthia  (at  anthesis) 
narrowly  campanulate,  mostly  2.5  mm  long  to 
the  torus,  moderately  to  sparsely  stellate  pubes- 
cent. Calyx  lobes  (on  fruiting  hypanthia)  persis- 
tent, glabrous,  erect,  broadly  semicircular  but 
±  truncate  apically,  entire  to  somewhat  erose, 
1  mm  long  and  about  2  mm  wide;  calyx  teeth 
persistent,  bluntly  subulate  or  knoblike,  ap- 
pressed  to  and  shorter  than  the  calyx  lobes  on 
mature  berries.  Petals  mostly  5,  but  4  in  some 
flowers,  reportedly  pink,  obovate,  entire,  round- 
ed to  irregularly  emarginate  at  apex,  3-5.5  mm 
long  and  2-4  mm  wide.  Stamens  10,  but  9  in  4- 
merous  flowers,  isomorphic;  filaments  straight, 


subulate,  thickened  and  somewhat  fleshy,  at 
least  basally,  2.5  mm  long;  anthers  2.5  mm  long 
and  mostly  1  mm  wide  distally,  yellow,  obovoid, 
border  of  apical  pore  ±  emarginate  ventrally  but 
truncate  dorsally;  connective  inconspicuous  and 
without  prolongations  or  appendages.  Ovary  in- 
ferior. Style  straight,  5  mm  long,  glandular-cil- 
iate  basally,  otherwise  glabrous;  stigma  subcap- 
itate.  Berry  ±  globose,  4.5-5.5  mm  long  to  the 
torus  and  5-5.5  mm  in  diameter.  Seeds  galei- 
form,  nitid,  smooth  to  obscurely  papillate,  0.5 
mm  long. 

TYPES. — Panama.  Bocas  del  Toro/Chiriqui  border:  Cerro 
Colorado  along  intersection  of  Bocas  Road  with  main  ridge 
road,  15.4  km  from  Chami  along  ridge  road,  elevation  1400- 
1700  m,  24  Oct.  1977,  Folsom  6143  (holotype:  CAS!;  isotype: 
MO). 

ADDITIONAL  SPECIMENS  EXAMINED. — Panama.  Chiriqui: 
Cerro  Colorado,  Bocas  Road,  Folsom  &  Collins  1748  (CAS, 
MO). 

DISTRIBUTION. — Known  only  from  Cerro 
Colorado  in  western  Panama  at  an  elevation  of 
1400-1700  m.  Flowering  and  fruiting  specimens 
have  been  collected  in  February  and  October. 

This  distinctive  species  is  characterized  by  a 
lax,  elongate  inflorescence,  truncate  calyx 
lobes,  nitid,  galeiform  seeds  and  cordate,  den- 
ticulate leaves  clothed  abaxially  with  sessile, 
stellate  trichomes.  In  the  few  collections  avail- 
able for  study,  the  inflorescence  is  consistently 
terminal  but  superficially  appears  to  diverge 
from  a  lateral  position  because  of  the  overtop- 
ping effect  created  by  lengthening  of  proximal 
axillary  shoots.  The  characteristic  number  of 
floral  parts  has  been  difficult  to  determine  on  the 
basis  of  material  at  hand.  The  inflorescence  on 
the  holotype  has  both  4-merous  flowers  with 
nine  stamens  and  5-merous  flowers  with  ten  sta- 
mens, but  the  significance  of  this  variation  can- 
not be  properly  assessed  without  a  more  exten- 
sive series  of  specimens. 

The  relationships  of  this  species  are  unclear. 
Assuming  sect.  Amblyarrhena  represents  a  nat- 
ural grouping,  it  is  tempting  to  suggest  that  M. 
coloradensis  diverged  from  the  Andean  stock 
that  gave  rise  to  M.  andreana  Cogn.  of  Colom- 
bia and  M.  gibba  Markgraf  of  Ecuador.  The  for- 
mer differs  from  M.  coloradensis  in  having  long- 
er leaves  (2-3.5  dm),  a  furfuraceous  pubescence, 
linear-oblong  to  subulate  bracteoles,  and  a  com- 
pact inflorescence  with  ultimate  units  consisting 
of  congested  glomerules.  Aside  from  striking 
differences  in  inflorescence  size  and  structure, 


ALMEDA:  NEW  SPECIES  OF  MICON1A 


307 


FIGURE  2.    Miconia  coloradensis  Almeda.  A,  habit,  x  ca.  1A;  B,  stamens,  ventral  view  (left)  and  3A  lateral  view  (right),  x  ca. 
6;  C,  mature  berry,  x3Vi,  D,  stellate  trichomes,  x  ca.  35.  E,  petal,  x7;  F,  seeds,  x!6.  (A-F  from  the  holotype.) 


the  new  species  approaches  M.  gibba  in  overall 
foliar  length  and  shape.  Petioles  of  the  latter  are 
shorter  (1.5-2  cm),  and  the  leaves  are  7-nerved, 
caducously  furfuraceous  below,  and  bigibbous 
dorsally  at  the  base. 

Miconia  concinna  Almeda,  nom.  nov. 

Topobea  micrantha  Pittier,  J.  Wash.  Acad.  Sci.  14:451.  1924. 
Nee  Miconia  micrantha  Cogn.  (Bull.  Torrey  Bot.  Club 
23:16.  18%)  nee  M.  micrantha  Pilger  (Verh.  Bot.  Ver. 


Brand.  47: 173.  1905;  M.  wittii  Ule,  nom.  nov.,  Notizbl.  Bot. 
Gart.  Berl.  6:367.  1915)  nee  M.  micrantha  Pittier  (Bol.  Soc. 
Venez.  Cienc.  Nat.  11:27.  1947;  M.  tabayensis  Wurdack, 
nom.  nov.,  Phytologia  21:359.  1971). 

Study  of  the  holotype  and  recently  collected 
material  of  this  entity  reveals  some  inaccuracies 
in  Pittier' s  incomplete  Latin  diagnosis.  It  seems 
appropriate,  therefore,  to  present  the  following 
emended  species  description. 


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Epiphytic  shrub  with  lax  arching  branches  to 

2  m  long.  Cauline  internodes  terete,  the  distal 
branches  glabrous  and  ±  vernicose  on  drying. 
Leaves  of  a  pair  isomorphic  to  anisomorphic, 
chartaceous,  ciliate-serrulate  (the  trichomes 
mostly  1-2  mm  long),  1.7-4.7  cm  long  and  1.4- 

3  cm  wide,  elliptic  to  elliptic-ovate  acuminate 
apically  and  acute  basally,  sparsely  pubescent 
to  glabrate  at  maturity  but  usually  lepidote  to 
brown-punctate  below,  sparsely  brown  furfura- 
ceous  to  glabrous  above,  the  pubescence  usually 
tardily  deciduous  and  imparting  a  brown-punc- 
tate appearance,  3-nerved  with  a  network  of  sec- 
ondaries mostly  1-2  mm  apart;  petioles  5-20  mm 
long,  1  mm  broad.  Inflorescence  a  pendant,  con- 
gested, trichotomously  branched  corymbiform 
panicle   1-1.5  cm  long  (shorter  than  foliage 
leaves  borne  at  the  node  initiating  the  inflores- 
cence) borne  on  a  stout  peduncle  2-5  mm  long; 
bracteoles  sessile,  foliaceous,  persisting  on  the 
infructescence,  oblanceolate  to  narrowly  spat- 
ulate,  3-6(-10)  mm  long,  1-2  mm  wide,  glabrous 
to  sparsely  furfuraceous.  Pedicels  terete,  gla- 
brous, mostly  0.5  mm  long.  Hypanthia  (at  an- 
thesis)  glabrous,  campanulate,  1-1.5  mm  long  to 
the  torus  and  1-1.5  mm  broad.  Calyx  lobes  (on 
fruiting  hypanthia)   ±   ascending,  persistent, 
semicircular  but  varying  to  rounded-deltoid,  en- 
tire to  minutely  lacerate,  1  mm  long  and  1.5  mm 
wide  at  base  between  sinuses;  calyx  teeth  per- 
sistent, triangular,  0.5  mm  long.  Petals  5,   ± 
erect  to  antrorsely  spreading,  glabrous,  ±  con- 
cave, white  but  sometimes  yellowish  on  drying, 
suborbicular,  entire,  2-2.5  mm  long  and  wide. 
Stamens    10,   isomorphic;  filaments  glabrous, 
subulate,  distally  geniculate,  2-2.5  mm  long;  an- 
thers 0.5  mm  long,  ±  cuneate,  broadly  flared 
and  terminated  by  a  ±  ovoid  ventrally  inclined 
pore;  connective  thickened  and  prolonged 
(0.5  x  0.5  mm)  below  thecae,  truncate  to  shal- 
lowly  bilobed  in  ventral  view,  dilated  dorsally 
into  a  blunt  projecting  appendage  in  profile  view. 
Ovary  ca.  4/5  inferior.  Style  straight,  glabrous, 
3  mm  long;  stigma  ±  clavate  to  subcapitate.  Ber- 
ry reportedly  black  at  maturity,  2-2.5  mm  long 
to  the  torus,  2.5-3  mm  in  diameter.  Seeds  ± 
pyriform,  smooth  and  nitid,  mostly  0.75  mm 
long. 

SPECIMENS  EXAMINED. — Panama.  Chiriqui:  humid  forests 
on  precipitous  slopes  of  Cerro  de  la  Horqueta,  Pittier  3276 
(US,  holotype  of  T.  micrantha);  s  slopes  of  Cerro  Horqueta 
N  of  Boquete,  Wilbur,  Teeri  &  Foster  13490  (CAS,  DUKE); 
Cerro  Pando,  on  continental  divide  and  Panama/Costa  Rica 


border,  ca.  16  km  NW  of  El  Hato  del  Volcan,  Mori  &  Bolten 
7291  (CAS),  Mori  &  Bolten  7301  (CAS). 

DISTRIBUTION. — Apparently  a  localized  cloud- 
forest  epiphyte  endemic  to  Chiriqui  province  in 
western  Panama  at  elevations  of  2000-2500  m. 
Flowering  and  fruiting  specimens  have  been  col- 
lected in  January,  March,  and  July. 

Miconia  concinna  is  recognized  by  its  lax, 
arching  branches,  adaxial  furfuraceous  indu- 
ment  on  juvenile  foliage,  large,  persistent,  folia- 
ceous floral  bracts,  and  pendant,  few-flowered 
corymbiform  inflorescences  that  are  markedly 
shorter  than  subtending  foliage  leaves. 

Pittier' s  (1924)  initial  placement  of  this  species 
in  Topobea  is  difficult  to  understand,  since  his 
discussion  makes  note  of  several  features  which 
made  this  decision  questionable.  In  choosing 
this  course,  Pittier  was  apparently  impressed  by 
the  presence  of  conspicuous  foliaceous  floral 
bracteoles  and  by  the  position  of  the  inflores- 
cence, which  he  erroneously  described  as  axil- 
lary. In  describing  this  species  as  a  tree,  it  also 
seems  likely  that  Pittier  mistook  its  habit  for  that 
of  its  host.  Label  Information  for  all  recently 
gathered  material  indicates  that  this  species  is 
an  epiphyte,  and  until  noted  otherwise,  it  seems 
advisable  to  accept  this  habital  description  as 
characteristic  of  the  species. 

On  the  basis  of  limited  material,  Standley 
(1938)  misinterpreted  M.  concinna  to  be  con- 
specific  with  the  Costa  Rican  endemic  described 
here  as  M.  longibracteata.  This  confusion  was 
compounded  when  he  referred  specimens  of 
these  two  taxa  to  M.  myrtillifolia  Naud.,  a 
species  of  Andean  Colombia  and  Venezuela 
which  differs  in  having  quadrate  branchlets,  a 
longer  (2-4  cm),  erect  panicle,  and  very  different 
oblong  anthers  with  diminutive  apical  pores  and 
unprolonged  connectives.  More  recently,  Glea- 
son  (1958)  included  M.  concinna  in  his  miscon- 
strued concept  of  M.  rubens  (Sw.)  Naud.  The 
latter,  a  dioecious  species,  is  known  only  from 
Jamaica  and  Venezuela  and  differs  most  notably 
by  its  elongate  multiflowered  inflorescence,  in- 
conspicuous bracteoles,  ±  swollen  nodes,  and 
different  anther  morphology. 

In  foliar  shape  and  floral  details,  M.  concinna 
and  M.  longibracteata  are  more  similar  to  each 
other  than  to  any  other  species  of  the  genus,  and 
there  is  little  doubt  that  they  were  derived  from 
common  ancestral  stock.  Available  collections 
suggest  that  these  two  species  are  allopatric, 
with  the  range  of  M.  concinna  lying  south  of 


ALMEDA:  NEW  SPECIES  OF  MICON1A 


309 


that  of  M.  longibracteata.  The  prevailingly  gla- 
brous leaves,  early-deciduous  floral  bracteoles, 
and  laxly  branched,  elongate  panicle  of  M.  lon- 
gibracteata serve  to  separate  these  species  most 
readily. 

Miconia  confertiflora  Almeda,  sp.  nov. 

(Figure  3) 

Sect.  Chaenopleura.  Frutex  epiphyticus  ca. 
2  m  altus.  Ramuli  sulcato-quadrangulati  sicut 
folia  inflorescentia  plerumque  glabri.  Petioli  3- 
16(-20)  mm;  lamina  1.7-7  x  1-3.4  cm  elliptica, 
elliptico-ovata  vel  obovata  apice  acuta  vel  acu- 
minata  basi  acuta,  3(-5)-nervata,  chartacea  et 
serrulata.  Panicula  corymbiformis,  pedunculo 
plus  minusve  1 .5  cm  longo;  flares  5-meri  bre viter 
(I  mm)  pedicellati,  bracteolis  I.5-2.5(-4)  mm 
longis  valde  caducis.  Hypanthium  (ad  torum) 
1.5  x  /  mm,  lobis  interioribus  0.5-1  mm  altis 
late  deltoideis  vel  rotundatis,  dentibus  exteriori- 
bus  acuminatis  0.5-1.5  mm  longis.  Petala  I  — 
1 .5  x  0.5-1  mm  ovata  apice  acuto  vel  paulo  un- 
cinato.  Stamina  isomorphica  glabra;  filamenta 
1—1.5  mm  longa;  antherarum  thecae  0.75-1  x 
0.25-0.50  mm  anguste  oblongae,  connectivum 
nee  prolongatum  nee  appendiculatum.  Stylus 
2  x  0.5  mm  glaber;  stigma  truncatum  non  ex- 
pans um. 

Epiphytic  shrub  to  2  m  tall,  distal  branches 
quadrangular  with  carinate  to  narrowly  alate  an- 
gles, entirely  glabrous  but  bearing  a  pair  of  ± 
pustulate  setiform  appendages  at  opposing  nodal 
faces.  Leaves  chartaceous,  glabrous,  1.7-7  cm 
long  and  1-3.4  cm  wide,  basally  entire  but  dis- 
tally  serrulate,  elliptic  but  sometimes  varying  to 
elliptic-ovate  or  obovate,  acute  to  acuminate 
apically  and  acute  basally,  3(-5)-nerved  with  a 
conspicuous  network  of  secondary  nerves,  dark 
green  above,  pale  green  and  occasionally  punc- 
tate below;  petioles  3-16(-20)  mm  long  and 
about  1  mm  broad.  Inflorescence  a  multiflow- 
ered  corymbiform  panicle  with  flowers  borne  in 
congested  terminal  glomerules;  rachis  glabrous, 
quadrangular,  mostly  less  than  1.5  cm  long; 
bracteoles  sessile,  glabrous,  early-deciduous, 
1.5-2.5(-4)  mm  long  and  0.5-1  mm  wide,  linear- 
subulate.  Pedicels  1  mm  long.  Hypanthia  (at  an- 
thesis)  campanulate,  glabrous,  1.5  mm  long  to 
the  torus.  Calyx  lobes  (on  fruiting  hypanthia) 
persistent,  erect,  broadly  deltoid  to  ±  rounded, 
entire  with  conspicuous  hyaline  margins,  0.5-1 
mm  long  and  1  mm  wide;  calyx  teeth  persistent, 


subulate  to  setiform,  equaling  or  commonly  ex- 
ceeding calyx  lobes  on  mature  berries.  Petals  5, 
glabrous,  erect  and  ±  concave  at  anthesis,  white 
but  tinged  with  red  externally,  narrowly  to 
broadly  ovate,  entire,  acute  to  bluntly  uncinate 
apically,  1-1.5  mm  long  and  0.5-1  mm  wide. 
Stamens  10,  isomorphic,  erect  to  slightly  in- 
curved at  anthesis;  filaments  white,  ±  translu- 
cent, subulate,  1-1.5  mm  long;  anthers  0.75-1 
mm  long  and  0.25-0.50  mm  wide  basally,  gla- 
brous, white,  narrowly  oblong  but  ±  obovoid  in 
profile  view,  distally  rounded  with  a  subtermi- 
nal,  oblong  ventrally  inclined  cleftlike  pore,  the 
margins  of  which  often  form  a  ±  elevated  hya- 
line border;  connective  thickened,  not  conspic- 
uously dilated  or  prolonged  dorsally  but  com- 
monly ±  prolonged  ventrally  below  thecae. 
Ovary  wholly  inferior.  Style  straight,  2  mm  long; 
stigma  truncate.  Berry  purple  at  maturity,  ±  glo- 
bose, 3-3.5  mm  long  to  the  torus  and  3.5  mm  in 
diameter.  Seeds  narrowly  ovoid  with  an  en- 
larged, ±  flattened  lateral  raphe,  densely  papil- 
late on  the  convex  surface,  mostly  2  mm  long 
and  1  mm  broad. 

TYPES. — Costa  Rica.  San  Jose:  about  18  km  N  of  San  Isidro 
de  Coronado  off  C.R.  #216  on  lower  w  slopes  of  Volcan  Ir- 
azii,  elevation  1700-1800  m,  5  July  1977,  Almeda  et  al.  2908 
(holotype:  CAS!;  isotypes:  CR!,  F!,  MO!,  US!). 

ADDITIONAL  SPECIMENS  EXAMINED. — Costa  Rica.  Here- 
dia:  slopes  NE  of  Cerro  Chompipe  about  16  km  NNE  of  San 
Rafael,  Wilbur,  Almeda  &  Daniel 22249  (CAS,  DUKE);  Cerro 
Zurqui,  NE  of  San  Isidro,  Standley  &  Valeria  50542  (US), 
Standley  &  Valeria  50644  (US);  saddle  area  between  Cerro 
Chompipe  and  SE  flank  of  Volcan  Barba  off  secondary  road 
N  of  C.R.  Hwy  #113  connecting  with  Calle  Gallito,  Baker, 
Utley  &  Utley  232  (CAS,  DUKE).  San  Jose:  5  km  NE  of  Cas- 
cajal,  Almeda  &  Nakai  3611  (CAS);  3-6  km  beyond  Las 
Nubes  in  vicinity  of  Cascajal,  Almeda  2636  (CAS);  about  7 
km  by  road  NE  of  Cascajal  and  14  km  NE  of  San  Isidro  de 
Coronado,  Wilbur  19816  (DUKE);  3-5  km  NE  of  Cascajal  in 
vicinity  of  Rio  Cascajal,  Wilbur  24420  (DUKE);  about  3  km 
NE  of  Cascajal  and  9  km  NE  of  San  Isidro  de  Coronado,  Wilbur 
19787  (DUKE). 

DISTRIBUTION. — A  local  cloud-forest  epi- 
phyte apparently  endemic  to  the  Cordillera  Cen- 
tral of  Costa  Rica  at  elevations  of  1600-2400  m. 
Flowering  and/or  fruiting  specimens  have  been 
collected  in  February,  March,  July,  October, 
and  December. 

Field  observations  and  label  information  of 
known  collections  indicate  that  this  species  is  an 
obligate  epiphyte.  Extirpation  of  moist  forests 
within  the  limited  range  of  this  taxon  poses  a 
real  threat  to  its  survival.  Fortunately,  some  in- 
dividuals persist  as  inhabitants  of  remnant  pas- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  vol.  42,  No.  10 


FIGURE  3.  Miconia  confertiflora  Almeda.  A,  habit,  xVi;  B,  representative  leaves,  upper  surface  (left)  and  lower  surface 
(right),  x%;  C,  fully  expanded  flower  showing  petals  and  stamens,  x9;  D,  petal,  x  16;  E,  stamens,  ventral  view  (left)  and 
lateral  view  (right),  x!3;  F,  mature  berries  with  persistent  calyx  lobes  and  calyx  teeth,  x3;  G,  seeds,  x6.  (A-G  from  Almeda 
etal.2908.) 


ture  trees  long  after  surrounding  vegetation  has 
been  cut  away. 

This  new  species  is  readily  separated  from 
congeners  by  its  quadrangular  branchlets,  dis- 
tally  serrulate  leaves,  congested  corymbiform 
inflorescence,  glabrous  hypanthia,  prominent 
calyx  teeth,  and  narrowly  ovoid  seeds  that  are 
densely  papillate  on  the  convex  side.  The  small, 


inconspicuous  anthers  are  also  noteworthy  in 
having  subterminal,  oblong  pores,  the  margins 
of  which  are  elevated  into  a  low,  continuous, 
hyaline  border.  This  feature  becomes  distorted 
with  pressing  and  drying  and  is  best  observed  in 
pickled  or  hydrated  material. 

Miconia  confertiflora  superficially  resembles 
M.  chionophylla  Naud.  of  sect.  Chaenopleura, 


ALMEDA:  NEW  SPECIES  OF  MICONIA 


311 


which  ranges  from  Andean  Colombia  to  Bolivia. 
The  latter  differs  in  its  procumbent  or  scandent 
habit,  puberulent  branchlets  and  petioles,  small- 
er leaves  (1-1.5  x  0.7-1.3  cm),  4-merous  flow- 
ers, and  capitate  stigma.  The  congested  inflo- 
rescence of  M.  confertiflora  is  also  reminiscent 
of  that  found  in  M.  parvifolia  Cogn.  (of  sect. 
Cremanium),  another  high-elevation  Colombian 
species  easily  separated  by  its  dwarf  shrubby 
habit  (2-3  dm  tall),  smaller,  revolute  leaves  (8- 
15  mm  long),  ovate,  apically  truncate  petals,  and 
copious  stellate  pubescence  on  distal  branchlets. 

Miconia  grandidentata  Almeda,  sp.  nov. 

Sect.  Chaenopleura.  Frutex  epiphyticus  ca. 
1  m  alt  us.  Ramuli  sulcato-quadrangulati  sicut 
folia  novella  primum  modice  vel  dense  pilis  stel- 
latis  induti  mox  glabrati.  Petioli  4-l5(-\9)  mm; 
lamina  3-6  x  1.8-3.5  cm  elliptica  vel  elliptico- 
ovata  apice  acuto  vel  acuminato  basi  acuta, 
3(-5)-nervata,  chart acea  obscure  distant erque 
serrulata.  Panicula  3-5.3  cm  longa  multiflora; 
flares  5-meri  breviter  (1-2  mm)  pedicellati,  brac- 
teolis  conspicuis  (l-)3~6  mm  longis  persistenti- 
bus.  Hypanthium  (ad  torum)  2-2.5  x  2  mm,  lob- 
is  interioribus  0.5  mm  altis  late  deltoideis  vel 
rotundatis,  dentibus  exterioribus  subulatis  2  mm 
longis.  Petala  1 .5-2  x  /  mm  plus  minusve  ovata 
apice  uncinato.  Stamina  isomorphica  glabra; 
filamenta  2  mm  longa;  antherarum  thecae  0.75- 
1  x  0.5  mm  oblongae,  rectae  vel  paulo  curvatae 
poro  ventraliter  inclinato,  connective  ad  basim 
dorsaliter  dente  hebeti  truncato  glabro  ornato. 
Stylus  2  x  0.5  mm  glaber;  stigma  truncatum 
non  expansum. 

Epiphytic  shrub  to  1  m  tall,  distal  branchlets 
moderately  to  densely  stellate,  quadrangular 
with  carinate  to  narrowly  alate  angles.  Older 
branches  somewhat  corky  on  drying,  cracking 
and  excorticating  in  age.  Leaves  chartaceous, 
entire  but  obscurely  serrulate  distally,  3-6  cm 
long  and  1.8-3.5  cm  wide,  elliptic  to  elliptic- 
ovate,  acute  to  acuminate  apically  and  acute  ba- 
sally,  glabrous  at  maturity  but  clothed  with 
brown  stellate  trichomes  when  young,  3(-5)- 
nerved  with  a  conspicuous  network  of  second- 
ary nerves  below;  petioles  4-15(-19)  mm  long 
and  1-2  mm  broad.  Inflorescence  an  erect,  ter- 
minal thyrse  mostly  3-5.3  cm  from  base  to  apex; 
the  rachis  prominently  quadrangular;  bracteoles 
sessile,  essentially  glabrous,  persistent,  gradu- 
ally reduced  in  size  upward,  (l-)3-6  mm  long 


and  0.5-2  mm  wide,  narrowly  lanceolate  to  sub- 
ulate and  ±  concave  adaxially.  Pedicels  1-1.5 
(-2)  mm  long  caducously  stellate  pubescent, 
each  pedicel  commonly  subtended  by  three 
bracteoles.  Hypanthia  (at  anthesis)  ±  globose, 
2-2.5  mm  long  to  the  torus,  beset  with  sessile 
stellate  trichomes  (these  early-deciduous  and 
generally  not  present  on  fruiting  hypanthia)  or 
persistent  only  as  remnant  arms  of  stellate  tri- 
chomes, which  superficially  resemble  puncti- 
form  glands.  Calyx  lobes  (on  fruiting  hypanthia) 
persistent,  sparsely  stellate  pubescent,  erect  to 
somewhat  incurved,  broadly  deltoid  to  ±  round- 
ed, entire  to  bluntly  undulate  with  hyaline  mar- 
gins, 0.5  mm  long  and  1  mm  wide;  calyx  teeth 
persistent,  subulate,  2  mm  long  and  1 .5  mm  wide 
at  base,  markedly  exceeding  calyx  lobes  on  ma- 
ture berries.  Petals  5,  glabrous,  erect  and  ±  con- 
cave at  anthesis,  white,  ±  ovate  in  outline,  en- 
tire, but  bluntly  uncinate  apically,  1.5-2  mm 
long  and  mostly  1  mm  wide.  Stamens  10,  iso- 
morphic,  erect  but  ±  incurved  and  exceeding 
the  style;  filaments  white  to  translucent,  subu- 
late, 2  mm  long;  anthers  0.75-1  mm  long  and 
mostly  0.5  mm  wide  basally,  glabrous,  white, 
oblong  to  rhomboid  in  profile  view,  but  broad- 
ened distally  to  an  oval  or  oblong,  ventrally  in- 
clined apical  pore;  connective  thickened  and 
prolonged  dorsally  at  the  base  into  a  deflexed  ± 
truncate  appendage.  Ovary  wholly  inferior. 
Style  straight,  2  mm  long;  stigma  truncate.  Berry 
pink  when  young  but  deep  purple  at  maturity, 
globose,  mostly  4  mm  long  to  the  torus  and  4 
mm  in  diameter.  Seeds  cuneate  and  conspicu- 
ously angled,  reddish  brown,  vernicose,  1.5- 
1.75  mm  long. 

TYPE. — Costa  Rica.  San  Jose:  About  5  km  NE  of  Cascajal, 
elevation  5400ft  [1646  m],  17  Feb.  1978,  Almeda  &  Nakai3627 
(holotype:  CAS!). 

ADDITIONAL  SPECIMENS  EXAMINED. — Costa  Rica.  Heredia: 
pastured  slopes  above  Rio  Para  Blanco  on  lower  slopes  of 
Cerro  Zurqui  about  7  km  NE  of  San  Josecito,  Wilbur  &  Luteyn 
18634  (DUKE).  San  Jose:  slopes  and  thickets  at  Alto  La  Pal- 
ma  about  15  km  in  a  straight  line  NE  of  San  Jose,  Wilbur  20340 
(CAS,  DUKE). 

DISTRIBUTION. — A  rare  cloud-forest  epiphyte 
known  only  from  the  south-facing  slopes  of  the 
Cordillera  Central  of  Costa  Rica  at  elevations  of 
1600-1800  m.  Flowering  material  has  been  col- 
lected in  February,  July,  and  December. 

Initial  study  of  this  species  led  me  to  interpret 
it  as  an  atypical  large-leafed  variant  of  M.  con- 
fertiflora, a  species  that  grows  in  the  same  gen- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  vol.  42,  No.  10 


eral  area.  Subsequent  field  observations  of  ad- 
ditional flowering  and  fruiting  material  indicate 
that  this  is  a  distinctive  taxon  worthy  of  specific 
rank.  Miconia  confertiflora  and  M.  grandiden- 
tata  share  an  epiphytic  habit,  quadrangular 
branchlets,  and  elliptic  to  elliptic-ovate,  distally 
serrulate  leaves  having  a  conspicuous  network 
of  secondary  nerves  below.  Miconia  grandiden- 
tata  differs  consistently  in  several  diagnostic 
characters.  It  lacks  the  nodal,  setiform  append- 
ages so  characteristic  of  distal  branchlets  in  M. 
confertiflora.  It  also  differs  in  having  stellate 
pubescence  on  branchlets  and  young  hypanthia, 
an  elongate  thyrsoid  panicle,  persistent  floral 
bracteoles,  longer  calyx  teeth,  and  angulate, 
vernicose  seeds  that  lack  a  well-defined  pro- 
longed lateral  raphe.  The  stamens  of  M.  gran- 
didentata  also  provide  several  distinguishing 
features.  Hydrated  anthers,  which  are  oblong  to 
rhomboid  in  profile  view,  are  broadened  distally 
to  an  oval  or  oblong,  ventrally  inclined  terminal 
pore,  and  the  connective  is  thickened  and  pro- 
longed dorsally  into  a  deflexed  caudiform  ap- 
pendage. 

Miconia  longibracteata  Almeda,  sp.  nov. 
(Figure  4) 

Sect.  Chaenopleura.  Frutex  1-3  m  altus.  Ra- 
muli  glabri  obscure  quadrangulati  demum  te- 
retes.  Petioli  7-!6(-27)  x  /  mm;  lamina  (1 .3-) 
2.5-4.2  x  1 .1-2.8  cm  chartacea  elliptica,  ellip- 
tico-obovata  aliquando  suborbicularia  apice 
acuta  vel  acuminata  basi  acuta  vel  obtusa,  tri- 
nervata  supra  primum  sparse  ferrugineo-furfu- 
racea  mox  glabrata,  subtus  primum  sparse  vel 
modice  lepidota  mox  glabrata.  Panicula  3.5-8 
cm  longa  multiflora;  flores  5-meri,  pedicelli  (1-) 
2.4  mm  longi,  bracteolis  4-1 1 (-18)  mm  longis 
oblongis  vel  anguste  spatulatis  usque  ad  an- 
thesim  persistentibus.  Hypanthium  (ad  torum) 
1-1.5  x  1 -1 .5  mm,  lobis  interioribus  1-1.5  mm 
altis  rotundatis  vel  deltoideis,  dentibus  exteriori- 
bus  acutis  0.5  mm  longis.  Petala  2-2.5  x  2  mm 
suborbicularia  glabra.  Stamina  isomorphica 
glabra;  filamenta  2.5  mm  longa;  antherarum 
thecae  0.5  x  0.25  mm  apice  late  biporosae, 
connective  sub  loculis  0.5  mm  prolongato  dor- 
saliter  ad  basim  (0.5  mm)  hebeti-tuberculato. 
Stylus  2-2.5  x  0.5  mm  glaber;  stigma  plus 
minusve  clavatum. 

Shrub  1-3  m  tall.  Cauline  internodes  glabrous, 
glossy  black  and  obscurely  quadrangular  when 


young,  becoming  brown  and  terete  with  age. 
Leaves  firmly  chartaceous,  ciliate-serrulate  (the 
trichomes  mostly  0.5-1  mm  long),  (1.3-)2.5-4.2 
cm  long  and  1.1-2.8  cm  wide,  elliptic,  elliptic- 
obovate  or  sometimes  varying  to  suborbicular, 
acuminate  to  acute  apically  and  acute  to  obtuse 
basally,  glabrous  at  maturity  but  caducously  lep- 
idote  below  and  glabrous  to  sparsely  brown  fur- 
furaceous  above  when  young,  3-nerved,  the  sec- 
ondaries conspicuous  below  and  mostly  2  mm 
apart;  petioles  7-16(-27)  mm  long  and  1  mm 
broad.  Inflorescence  an  erect,  laxly  branched, 
elongate  panicle  3.5-8  cm  long,  exceeding  fo- 
liage leaves  borne  at  the  node  initiating  the  in- 
florescence; rachis  glabrous,  quadrate  to  ± 
rounded;  bracteoles  sessile,  foliaceous,  decidu- 
ous following  anthesis  and  mostly  absent  on  the 
infructescence,  linear-oblong  to  narrowly  spat- 
ulate,  ±  concave  to  navicular,  4-ll(-18)  mm 
long,  0.5-3  mm  wide,  glabrous  above  and  below. 
Pedicels  terete,  glabrous,  (l-)2-4  mm  long.  Hy- 
panthia (at  anthesis)  glabrous,  campanulate,  1- 
1.5  mm  long  to  the  torus  and  1-1.5  mm  broad 
distally.  Calyx  lobes  (on  fruiting  hypanthia) 
erect  to  ascending,  persistent,  semicircular  but 
varying  to  ±  deltoid,  entire,  1-1.5  mm  long  and 
1-1.5  mm  wide  basally  between  sinuses;  calyx 
teeth  persistent,  triangular,  0.5  mm  long.  Petals 
5,  antrorsely  spreading,  glabrous,  ±  concave, 
white  to  yellowish  white,  suborbicular,  entire, 
2-2.5  mm  long,  2  mm  wide.  Stamens  10,  iso- 
morphic;  filaments  glabrous,  subulate,  white, 
distally  incurved,  2.5  mm  long;  anthers  about 
0.5  mm  long  or  less,  0.25  mm  wide  distally, 
white,  ±  cuneate,  flared  distally  and  terminated 
.by  a  broad,  ±  ovoid,  ventrally  inclined  pore; 
connective  markedly  thickened  and  prolonged 
(0.5  x  0.5  mm)  below  thecae,  dilated  dorsally 
into  a  blunt  knobby  protuberance.  Ovary  totally 
inferior.  Style  straight,  glabrous,  2-2.5  mm  long; 
stigma  ±  clavate.  Berry  deep  purple  to  purple- 
black  at  maturity,  3.5-5  mm  long  to  the  torus, 
3-4(-5)  mm  in  diameter.  Seeds  ±  pyriform,  ob- 
scurely muriculate,  mostly  1  mm  long. 

TYPES. — Costa  Rica.  Alajuela:  wooded  slopes  of  Volcan  Poas 
about  12  km  w  of  Varablanca,  elevation  ca.  2400  m,  21  Jan. 
1968,  Wilbur  &  Stone  9845  (holotype:  DUKE!;  isotypes: 
CAS!,  US!). 

ADDITIONAL  SPECIMENS  EXAMINED. — Costa  Rfca.  Alajue- 
la: Volcan  Poas,  forest  between  crater  and  cold  lake,  Davidse 
&  Pohl  1169  (US);  Volcan  Poas,  cloud  forest  along  road  2.5- 
3.5  miles  [4.0-5.6  km]  w  of  Poasito,  Webster,  Miller  &  Miller 
12242  (DUKE,  US);  SE  slope  of  Volcan  Poas,  Hatheway  1388 


ALMEDA:  NEW  SPECIES  OF  M1CONIA 


313 


FIGURE  4.  Miconia  longibracteata  Almeda.  A,  habit,  x  ca.  %;  B,  seeds,  x!4;  C,  petal,  x9;  D,  representative  leaves,  upper 
surface  (left)  and  lower  surface  (right),  xl;  E,  mature  berry,  x  ca.  4;  F,  stamens,  lateral  view  (left)  and  %  ventral  view  (right), 
xlO:  G,  stamens,  dorsal  view  (left)  and  ventral  view  of  anther  and  prolonged  connective  (right),  xlO.  (A  &  D  from  Schnell 
727;  B,  C,  E-G  from  Wilbur  &  Stone  9845.) 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  vol.  42,  No.  10 


(DS,  US).  Cartago:  El  Empalme,  Schnell  727  (US— 2  sheets); 
Talamanca  Range,  Panamerican  Highway,  Carlson  3611  (US); 
El  Canon  (Carretera  Panamericana),  O.  Jimenez  42  (US);  La 
Chonta,  Schnell  725  (US);  Panamerican  Highway  about  24  km 
w  of  Villa  Mills,  Wilbur  &  Stone  8800  (DUKE);  s  of  El  Em- 
palme, A.  Jimenez  1979  (US).  Cartago/San  Jose  border:  6  km 
NW  of  Dos  Amigos,  Wilbur  &  Luteyn  18321  (DUKE);  5  km 
NW  of  Ojo  de  Agua  or  19  km  SE  of  El  Empalme,  Wilbur  28674 
(DUKE);  5  km  SE  of  Trinidad  along  Carretera  Interamericana, 
Wilbur  27780  (DUKE). 

DISTRIBUTION. — Endemic  to  Costa  Rica  where 
it  is  presently  known  from  the  slopes  of  Volcan 
Poas  and  the  Cordillera  de  Talamanca  at  eleva- 
tions of  2000-3000  m.  Available  collections  in- 
dicate that  flowering  and  fruiting  occurs  sporad- 
ically throughout  the  year. 

Among  Costa  Rican  taxa  of  Miconia,  this 
species  is  recognized  by  the  combination  of 
large  foliaceous  floral  bracteoles,  prevailingly 
glabrous  foliage,  suborbicular  petals,  and  pecu- 
liar androecial  morphology  (see  Fig.  4F).  Sta- 
minal  posture  and  morphology  are  best  observed 
in  pickled  or  boiled  material  hydrated  with  the 
aid  of  a  wetting  agent.  Materials  so  treated  show 
short,  compressed,  anther  thecae  terminating  in 
a  broad  apical  pore;  the  connective  which  is 
conspicuously  thickened  and  prolonged  below 
the  thecae  is  narrowly  triangular  in  ventral  view, 
knobby  and  somewhat  angulate  in  profile  view, 
and  ±  horseshoe  shaped  in  dorsal  view.  In  many 
respects  the  elaborate  connective  is  reminiscent 
of  a  pedestal  providing  anchorage  and  support 
for  the  anther.  This  distinctive  anther  morphol- 
ogy appears  to  represent  an  extreme  in  the  evo- 
lutionary line  that  has  led  to  great  reduction  in 
anther  size.  The  very  broad  apical  pores  and 


pronounced  geniculation  of  the  filaments  consis- 
tently bring  anthers  to  an  incurved  position 
within  the  flower  and  may  contribute  to  preva- 
lent self-pollination. 

In  size  and  shape  of  floral  bracteoles  and  an- 
thers, M.  longibracteata  is  most  similar  to  M. 
concinna  (also  treated  herein).  The  characters 
distinguishing  these  species  are  enumerated  in 
the  discussion  under  M.  concinna.  Miconia  lon- 
gibracteata also  resembles  M.  superposita  Wur- 
dack  in  vegetative  aspect.  The  latter,  a  Colom- 
bian species,  differs  in  having  squamulose 
pubescence  on  juvenile  branchlets,  shorter  brac- 
teoles (3.7  x  0.8  mm),  and  4-celled  anthers  with 
the  dorsal  loculus  of  each  theca  overlapping  but 
distal  to  the  ventral  one. 

ACKNOWLEDGMENTS 

I  thank  Terry  Bell  for  preparing  the  line  draw- 
ings, Dr.  J.  J.  Wurdack  for  review  of  the  manu- 
script, the  Museo  Nacional  de  Costa  Rica  and 
the  Organization  for  Tropical  Studies  for  logis- 
tical support,  and  curators  of  the  following  her- 
baria for  special  loans  or  use  of  their  facilities: 
CR,  DUKE,  F,  MO,  US.  This  study  was  sup- 
ported by  U.S.  National  Science  Foundation 
Grants  DEB  76-83040  and  DEB  78-25620. 

LITERATURE  CITED 

GLEASON,  H.  A.    1958.   Melastomataceae.  Flora  of  Panama. 

Ann.  Missouri  Hot.  Gard.  45:203-304. 
PITTIER,  H.    1924.    New  or  little  known  Melastomataceae 

from  Venezuela  and  Panama.  II.  J.  Wash.  Acad.  Sci. 

14:447_451. 
STANDLEY,  P.  C.   1938.  Flora  of  Costa  Rica.  Field  Mus.  Nat. 

Hist.,  Bot.  Ser.  18(3):783-1133. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  941 18 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  11,  pp.  315-322,  5  figs. 


June  24,  1981 


STUDIES  ON  CAVE  HARVESTMEN  OF  THE  CENTRAL  SIERRA 

NEVADA  WITH  DESCRIPTIONS  OF 

NEW  SPECIES  OF  BANKSULA 

By 
Thomas  S.  Briggs 

Research  Associate,  Department  of  Entomology,  California  Academy  of  Sciences,  San  Francisco,  California  94118 

and 
Darrell  Ubick 

Biology  Department,  San  Jose  State  University,  San  Jose,  California  95114 


ABSTRACT:  New  ecological  and  biogeographic  information  on  Sierra  Nevada  cave  harvestmen  in  Banksula 
was  obtained  while  environmental  impact  and  mitigation  work  was  being  done  for  the  Army  Corps  of  Engineers 
New  Melones  Dam  project.  Isolation  appears  to  be  the  principal  factor  leading  to  speciation  in  Banksula,  but 
the  distribution  of  species  in  the  vicinity  of  the  New  Melones  Reservoir  is  not  readily  explained.  Four  new 
species  of  Banksula  are  described:  It.  rudolphi,  B.  martinorum,  B.  grubbsi,  and  It.  elliotti. 


INTRODUCTION 

Troglobitic  organisms  are  scarce  in  California 
caves,  possibly  due  to  the  relatively  small  size 
and  geologic  youth  of  these  habitats.  Laniatorid 
harvestmen  of  the  genus  Banksula  are  distinc- 
tive because  they  are  relatively  abundant  obli- 
gate cavernicoles  of  the  Calaveras  Formation  of 
the  Sierra  Nevada.  Intensive  collecting  by  bio- 
speleological  investigators  contracted  by  the 
Army  Corps  of  Engineers,  the  Fish  and  Wildlife 
Service,  and  the  Office  of  Endangered  Species 
has  yielded  numerous  new  records  and  four  new 
species  of  Banksula.  Project  teams  worked  pri- 
marily in  the  vicinity  of  the  New  Melones  Res- 
ervoir site  on  the  Stanislaus  River,  Calaveras 
and  Tuolumne  counties,  where  several  caves  are 
threatened  by  completion  of  the  New  Melones 
Dam.  Biological  surveys  were  begun  in  May 


1975  when  a  mine  tunnel  in  limestone  was  se- 
lected for  transplanting  biota,  including  Bank- 
sula grahami  and  B.  melones,  from  McLean's 
Cave,  the  largest  of  the  threatened  caves.  As 
additional  workers  transplanted  animal  and 
plant  life  from  McLean's  Cave  and  studied  other 
nearby  caves,  some  distributional,  behavioral, 
and  ecological  information  were  obtained  which 
allow  us  to  present  some  biogeographic  discus- 
sion of  Banksula. 

Briggs  (1974)  reviewed  and  expanded  the  ge- 
nus Banksula  and  described  four  species  with 
functional  eyes.  He  showed  that  the  single  pre- 
viously known  species,  Banksula  calif ornica 
(Banks),  lacked  corneas  and  retinae.  His  con- 
tention that  all  Banksula  are  confined  to  caves 
is  supported  by  all  subsequent  investigations. 
Only  Banksula  melones,  which  has  relatively 


[315] 


316 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  11 


well-developed  eyes,  has  been  collected  near  or  nose  tubercle  on  patella;  retinae  incom- 

at  the  cave  twilight  zone.  Sitalcina  sierra  Briggs  plete  or  absent;  corneas  present  6 

and  Horn  is  the  only  laniatorid  harvestman 

,.        ,  .                   ,    ,  .           ,.  5a.  Submargmal  row  of  tubercles  on  eighth 

found  in  epigean  habitats  adjacent  to  cave  lo- 

..,.  tergite  and  last  sternite    . 

cahties.  .                " 

__  B.  ruaolpni  new  species 

METHODS  AND  DEPOSITION  5b.  No  submarginal  rows  of  tubercles  on  ter- 

gites  or  sternites 

The  structures  measured  and  morphological  „ 

B.  martmorum  new  species 

terminology  used  in  this  paper  are  generally  6a    Eye  tuberde  a  rounded  CQne;  small  spur 

those  descnbed  by  Bnggs  (1968)  for  laniatorid  ectal  to  proximal  ventral  spine  on  palpal 

harvestmen,  with  one  modification  that  appears  femur;  incomplete  retinae  usuany  pres. 

in  this  paper  and  in  Briggs  (1974):  we  substitute  ent B  tuo[umne  Briggs 

"second  endites"  for  that  portion  of  the  second  6b    Eye  tubercle  obliquely  truncate  in  lateral 

coxae  previously  referred  to  as  the  maxillary  view;  no  spur  ectal  to  proximal  ventral 

processes.  Another  structure,  the  labial  process,  spine  on  palpal  femur;  retinae  absent 

first  described  by  Briggs  (1971),  is  located  be-  B  gm^si  new  species 

tween  the  second  endites. 

Unless  otherwise  noted,  all  specimens  record-  7a-  First  legs  longer  than  4  mm;  retinal  di- 
ed in  this  paper  were  preserved  in  ethyl  alcohol  ameter  greater  than  0.03  mm;  aedeagus 
and  are  deposited  in  the  collection  of  the  Cali-  and  enclosing  sheath  fold  anterior  to  re- 
fornia  Academy  of  Sciences.  curved  prongs  of  dorsal  plate  _ 

B.  melones  Briggs 

Key  to  the  Species  of  Banksula  7b.  First  legs  shorter  than  4  mm;  retinal  di- 

la.  Operculum  small,  held  almost  entirely  be-  ameter  less  than  0.03  mm  or  missing;  ae- 

tween  mesal  margins  of  fourth  coxae;  deagus  and  enclosing  sheath  held  within 

males  with  apex  of  aedeagus  not  enclosed  bifurcate  dorsal  plate  __.                            _   8 

in  sheath,  velum  on  dorsal  plate  smooth  8a    Eye  tubercle  and  elevated  area  behind 

(californica  group)  without  tubercles;  retinae  present 

Ib.  Operculum  large,  with  posterior  margin  B  grahami  Briggs 

well  behind  mesal  extreme  of  posterior  8b    Eye  tubercle  and  eievated  area  behind  tu- 

margin  of  fourth  coxae;  males  with  apex  berculate;  retinae  reduced  or  missing  .... 

of  aedeagus  enclosed  in  sheath,  velum  on  B  elliotti  new  species 
dorsal  plate  in  transverse  folds  (melones 

group) ___                                                          7  Banksula  rudolphi  new  species 

2a.  Retinae  entire;  eye  tubercle  obliquely  (Figure  2) 

truncate  in  lateral  view  ____  B.  galilei  Briggs  HOLOTYPE. — Male.   Body  length   1.67  mm; 

2b.  Retinae  absent  or  incomplete;  if  retinae  scute  length  1.40  mm;  scute  width  1.14  mm;  eye 

present,  eye  tubercle  a  rounded  cone   ..    3  tubercle  length  0.23  mm;  eye  tubercle  width  0.28 

mm;  operculum  length  0.18  mm;  operculum 

3a.  Small  spines  or  tubercles  between  pnn-  . ,  ,    ' ., 

.  _  width  0.21  mm. 
cipal  dorsal  spines  on  palpal  femur;  cor- 

Palp:  trochanter  0.19  mm,  femur  0.79  mm, 

neas  absent  B.  cahformca  (Banks)  .  „    „  . .           ....    „  ,c                      „  .. 

_.                            .  patella  0.44  mm,  tibia  0.65  mm,  tarsus  0.44  mm. 

3b.  No  small  spines  or  tubercles  between  T       TT          ,            _    n           ,, 

.     .                 .       .                        .  _  Leg  II:  trochanter  0.19  mm,  femur  1.42  mm, 

principal  dorsal  spines  on  palpal  femur;  .  .              ...    ,  _ ,                              .  __ 

patella  0.44  mm,  tibia  1.26  mm,  metatarsus  1.02 

corneas  present  or  absent  4  ,      ' 

mm,  tarsus  1.58  mm. 

4a.  Proximal  ventral  spinose  tubercle  on  pal-  Scute  with  segmentation  delineated  by  tuber- 
pal  tibia  equal  to  ventral  spinose  tubercle  cles,  eye  tubercle  and  area  behind  tuberculate. 
on  patella;  retinae  absent;  corneas  pres-  Eye  tubercle  a  rounded  cone  without  retinae 
ent  or  absent  5  (some  individuals  also  without  corneas).  Ter- 

4b.  Proximal  ventral  spinose  tubercle  on  pal-  gites  with  row  of  tubercles  at  margin,  eighth  ter- 

pal  tibia  always  smaller  than  ventral  spi-  gite  also  with  medial  row  of  tubercles.  Posterior 


BRIGGS  AND  UBICK:  CAVE  HARVESTMEN 


317 


•  californica 

A  galilei 

•  grahami 

•  me/ones 

•  tuo/umne 
O  eliiotti 

A  grubbsi 
O  martini 
O  rudolphi 


FIGURE  1.     Map  showing  counties  in  the  central  Sierra  Nevada  where  Banksula  have  been  collected.  The  inset  shows  the 
region  along  the  Stanislaus  River  where  environmental  impact  and  mitigation  work  on  Banksula  has  been  concentrated. 


sternite  with  two  rows  of  tubercles.  Anal  plate 
with  medial  tubercles.  Second  endites  with  con- 
cave ectal  margin.  Labial  processes  rounded. 
Operculum  with  narrow,  rounded  anterior;  pos- 
terior margin  adjacent  to  mesal  extreme  of  pos- 
terior margin  of  fourth  coxae. 

Palpal  femur  with  six  prominent  proximal  dor- 
sal spines  (some  individuals  with  only  five 
spines)  and  one  small  distal  dorsal  spine;  no  se- 
tae or  tubercles  between  dorsal  spines;  ectal 
spur  at  base  of  first  ventral  spine  small  and 
rounded;  venter  with  numerous  small  tubercles. 
Proximal  ventral  tubercle  on  palpal  tibia  re- 
duced, spine  from  tubercle  absent.  Ventral  spine 
on  palpal  patella  arising  from  a  reduced  tubercle. 

Tarsal  formula  4-6-5-6. 

Body  concolorous  yellow-orange. 

Penis  typical  of  californica  group  (see  Briggs 
1974:11-12). 

ALLOTYPE. — Female.  Slightly  smaller  than 


holotype.  Proximal  ventral  spinose  tubercle  on 
palpal  tibia  present,  equal  in  size  to  robust  ven- 
tral spinose  tubercle  on  palpal  patella.  Opercu- 
lum with  truncate  apex  (rounded  in  some  indi- 
viduals); posterior  margin  behind  mesal  extreme 
of  posterior  margin  of  fourth  coxae. 

TYPE-SPECIMENS.— Holotype,  allotype,  21  paratypes  (9  3, 
99,3  juv.):  Chrome  Cave,  near  Jackson,  Amador  County, 
California,  5  Apr.  1979,  D.  C.  Rudolph,  S.  Winterath,  and  B. 
Martin.  Paratypes,  2  6,  2  9,  3  juv.:  same  locality,  21  Sep. 
1980,  T.  S.  Briggs  and  D.  Ubick.  Paratype,  1  6:  same  locality, 
24  Jan.  1981,  T.  S.  Briggs  and  D.  Ubick. 

ETYMOLOGY. — This  species  is  named  for 
biospeleologist  D.  Craig  Rudolph. 

ECOLOGICAL  NOTES. — Chrome  Cave  is  a  small 
cave  situated  in  about  0.02  km2  of  limestone. 
The  outcrop  is  surrounded  by  serpentine  which 
also  forms  much  of  the  cave's  walls.  The  hu- 
midity is  relatively  high  and  a  mean  temperature 
of  18  C  was  recorded  on  24  January  1981.  Bank- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 1 


FIGURE  2.    Banksula  rudolphi  Briggs  and  Ubick,  new  species,  a.  Lateral  view  of  body  and  left  palpus  of  male  holotype;  b. 
Lateral  view  of  left  palpus  of  female  allotype. 


sula  rudolphi  is  found  throughout  the  upper  re- 
gions of  the  cave,  predominantly  on  the  under- 
sides of  rocks.  Associated  with  Banksula  is  a 
rich  assortment  of  predators,  of  which  spiders 
(Araneae)  are  most  abundant.  We  recorded  the 
following  species:  Archoleptoneta  schusteri 
Gertsch,  Liocranoides  sp.,  Trogloneta  para- 
doxa  Gertsch,  and  Usofila  (Tele ma)  sp.  Of  the 
other  cavernicoles  encountered,  the  most  inter- 
esting were  specimens  of  Prokoenenia  sp.  (Pal- 
pigradida). 

Banksula  martinorum  new  species 

(Figure  3) 

HOLOTYPE. — Male.  Body  length  1.75  mm; 
scute  length  1.49  mm;  scute  width  1.23  mm;  eye 
tubercle  length  0.26  mm;  eye  tubercle  width  0.33 
mm;  operculum  length  0.18  mm;  operculum 
width  0.19  mm. 

Palp:  trochanter  0.19  mm,  femur  0.88  mm, 
patella  0.53  mm,  tibia  0.65  mm,  tarsus  0.42  mm. 

Leg  II:  trochanter  0.21  mm,  femur  1.81  mm, 
patella  0.39  mm,  tibia  1.67  mm,  metatarsus  1.23 
mm,  tarsus  2.37  mm. 

Scute  with  segmentation  delineated  by  small 
tubercles;  eye  tubercle  and  area  behind  slightly 
tuberculate.  Eye  tubercle  subconical,  without 


retinae  or  corneas.  Tergites  with  submarginal 
row  of  tubercles.  Second  endites  large,  ectal 
margin  slightly  concave.  Labial  processes  with 
acute  anterior  margin.  Operculum  with  rounded 
anterior,  posterior  margin  slightly  behind  mesal 
extreme  of  posterior  margin  of  fourth  coxae. 
First  coxae  with  two  prominent  spinose  tuber- 
cles. Anal  plate  smooth. 

Palpal  femur  with  five  prominent  proximal 
dorsal  spines  and  two  slightly  smaller  distal  dor- 
sal spines;  no  setae  or  tubercles  between  dorsal 
spines;  ectal  spur  at  base  adjacent  to  second 
ventral  spine.  Proximal  ventral  spinose  tubercle 
on  palpal  tibia  equal  to  ventral  spinose  tubercle 
on  patella. 

Tarsal  formula  4-6-5-6. 

Body  concolorous  pale  yellow. 

Penis  typical  of  calif  arnica  group. 

ALLOTYPE. — Female.  Similar  to  male. 

TYPE-SPECIMENS. — Holotype.  allotype,  3  paratypes  (1  9,  2 
juv.):  Heater  Cave,  8  km  N  Columbia,  Calaveras  County,  Cal- 
ifornia, 15  Mar.  1979,  D.  C.  Rudolph,  B.  Martin,  and  S.  Win- 
terath. 

ETYMOLOGY. — This  species  is  named  for 
arachnologist  Barbara  Martin  and  for  Army 
Corps  of  Engineers  environmental  planner  Rob- 
ert Martin. 


BRIGGS  AND  UBICK:  CAVE  HARVESTMEN 


319 


FIGURE  3.    Banksula  martinorum  Briggs  and  Ubick,  new  species.  Lateral  view  of  body  and  left  palpus  of  male  holotype. 


Banksula  grubbsi  new  species 

(Figure  4) 

HOLOTYPE. — Male.  Body  length  1.49  mm; 
scute  length  1.26  mm;  scute  width  1.23  mm;  eye 
tubercle  length  0.25  mm;  eye  tubercle  width  0.35 
mm;  operculum  length  0.18  mm;  operculum 
width  0.21  mm. 

Palp:  trochanter  0.19  mm,  femur  0.70  mm, 
patella  0.39  mm,  tibia  0.53  mm,  tarsus  0.65  mm. 

Leg  II:  trochanter  0.18  mm,  femur  1.49  mm, 
patella  0.44  mm,  tibia  1.23  mm,  metatarsus  0.96 
mm,  tarsus  1.49  mm. 

Scute  with  segmentation  delineated  by  small 
tubercles,  area  behind  eye  tubercle  tuberculate. 
Eye  tubercle  obliquely  truncate,  declining  pos- 
teriorly; entire  surface  tuberculate.  Eyes  with 
small  corneas  and  no  retinae.  Tergites  with  sub- 
marginal  row  of  tubercles.  Second  endites  se- 
tose, with  acute  ectal  invagination  behind  mid- 
point. Labial  processes  large,  rounded.  Small 
operculum  with  rounded  anterior,  margin  adja- 
cent to  fourth  coxae,  posterior  margin  slightly 
behind  mesal  extreme  of  posterior  margin  of 
fourth  coxae. 

Chelicerae  setose,  only  slightly  tuberculate. 

Palpal  femur  with  four  prominent  proximal 
dorsal  spines  and  three  reduced  distal  dorsal 
spines;  no  setae  or  tubercles  between  proximal 
dorsal  spines;  ectal  spur  at  base  replaced  by  a 
broad,  low  tubercle;  ventral  margin  straight. 
Palpal  tibia  without  significant  proximal  ventral 
spine. 

Tarsal  formula  4-6-5-6. 

Body  concolorous  light  yellow. 

Penis  typical  of  calif  arnica  group,  aedeagal 
velum  held  in  smooth  dorsal  plate. 


TYPE-SPECIMEN. — Holotype:  Black  Chasm  Cave,  near  Vol- 
cano, Amador  County,  California,  19  Feb.  1978,  A.  G. 
Grubbs. 

FEMALE. — Unknown. 

ETYMOLOGY. — This  species  is  named  for 
biospeleologist  Andrew  G.  Grubbs. 

Banksula  elliotti  new  species 

(Figure  5) 

HOLOTYPE. — Male.  Body  length  1.61  mm; 
scute  length  1.23  mm;  scute  width  1.28  mm;  eye 
tubercle  length  0.26  mm;  eye  tubercle  width  0.35 
mm;  operculum  length  0.30  mm;  operculum 
width  0.32  mm. 

Palp:  trochanter  0.23  mm,  femur  0.63  mm, 
patella  0.39  mm,  tibia  0.53  mm,  tarsus  0.39  mm. 

Leg  II:  trochanter  0.16  mm,  femur  1.14  mm, 
patella  0.35  mm,  tibia  0.96  mm,  metatarsus  0.72 
mm,  tarsus  1.14  mm. 

Scute  with  segmentation  delineated  by  tuber- 
cles, eye  tubercle  and  area  behind  tuberculate. 
Eye  tubercle  rounded,  slightly  conical,  with 
small  corneas  and  without  retinae.  Tergites  with 
row  of  tubercles  at  margin.  Second  endites  se- 
tose, mesally  broad  and  rounded;  labial  pro- 
cesses spatulate.  Operculum  large,  extending 
posterior  to  hind  coxae.  Hind  sternite  and  anal 
plate  with  row  of  tubercles.  All  coxae  tubercu- 
late. 

Chelicerae  with  strongly  tuberculate  anterior 
margin. 

Palpal  femur  with  six  prominent  proximal  dor- 
sal spines  and  four  reduced  distal  dorsal  spines; 
no  setae  or  tubercles  between  proximal  dorsal 
spines;  ectal  spur  well  developed  at  base  of 


320 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  11 


FIGURE  4.    Banksula  grubbsi  Briggs  and  Ubick,  new  species.  Lateral  view  of  body  and  left  palpus  of  male  holotype. 


proximal  ventral  spine,  ventral  surface  tuber- 
culate. 

Tarsal  formula  4-6-5-6. 

Body  concolorous  yellow-orange. 

Penis  with  folds  at  base  of  dorsal  plate;  apex 
of  aedeagus  enclosed  in  rounded  sheath  and  held 
within  distal  bifurcation  of  dorsal  plate. 

ALLOTYPE. — Female.  Similar  to  male. 

TYPE-SPECIMENS. — Holotype,  allotype,  4  paratypes  (2  $ ,  1 
2 ,  1  juv.):  Pinnacle  Point  Cave,  near  Camp  Nine  Power 
House,  9.6  km  N  Columbia,  Tuolumne  County,  California,  8 
Feb.  1979,  D.  C.  Rudolph,  B.  Martin,  and  S.  Winterath.  Para- 
types,  2  <5,  2  juv.:  same  locality,  20  Jan.  1978,  W.  Elliott,  A. 
Grubbs,  and  S.  Winterath.  Paratypes,  2  3 ,  1  9 :  same  locality, 
1  May  1977,  A.  Grubbs,  N.  Boice,  M.  McEachern  and  J. 
Davis.  Paratypes,  1  9:  Digger  Pine  Cave,  6.5  km  N  Columbia, 
Calaveras  County,  22  Mar.  1979,  D.  Rudolph,  B.  Martin,  S. 
Winterath,  W.  Elliott,  and  J.  Reddell.  Paratypes,  3  3,5  2: 
Rabbit  Hole  Cave,  6.5  km  N  Columbia,  Calaveras  County,  22 
Mar.  1979,  D.  Rudolph,  B.  Martin,  and  S.  Winterath.  Para- 
types, 1  6,  1  2:  Grapevine  Gulch  Cave,  6.5  km  N  Columbia, 
Calaveras  County,  22  Nov.  1979,  D.  Rudolph,  B.  Martin,  and 
S.  Winterath.  Paratypes,  1  d,  4  2:  same  locality,  27  Apr. 
1977,  B.  Hawson,  A.  Grubbs,  J.  Munthe,  and  M.  McEachern. 

ETYMOLOGY. — This  species  is  named  for 
biospeleologist  William  R.  Elliott. 

VARIATIONS. — Specimens  taken  in  Pinnacle 
Point  Cave  are  without  retinae,  although  half  of 
the  specimens  taken  in  the  remaining  caves  have 
retinal  pigment  in  their  eyes.  The  specimen  from 
Digger  Pine  Cave  has  a  relatively  more  conical 
eye  tubercle.  Some  specimens  have  only  three 
distal  dorsal  spines  on  the  palpal  femur. 

Banksula  grahami  Briggs 

Banksula  grahami  BRIGGS  1974:7. 

RECORDS. — Calaveras  County:  Moaning  Cave,  near  Valle- 
citos,  22  Aug.  1963,  22  Dec.  1968,  6  Dec.  1977,  R.  Graham, 
T.  Briggs,  W.  Elliott,  A.  Grubbs,  and  S.  Winterath;  Linda's 
Cave,  6  km  w  Columbia,  16  May  1977,  A.  Grubbs,  N.  Boice, 
and  D.  Broussard;  Carlow's  Cave,  6  km  w  Columbia,  16  May 


1977,  A.  Grubbs,  N.  Boice,  and  D.   Broussard.   Tuolumne 
County:  Experimental  Mine  Cave,  3  km  N  Columbia,  25  Jun. 
1975,  R.  Lem;  Mine  tunnel  along  road  to  Experimental  Mine, 
2.5  km  N  Columbia,  25  Jun.   1975,  T.  Briggs  (identification 
only);  Snell's  Cave,  3  km  N  Columbia,  26  Feb.  1978,  14  Feb. 

1978,  2  Apr.  1979,  S.  Winterath,  D.  Rudolph,  and  J.  Reddell; 
Crystal  Palace  Cave,  5  km  N  Columbia,  4  Nov.  1967,  21  Dec. 
1977,  4  Feb.  1979,  25  Mar.  1979,  T.  Briggs,  V.  Lee,  D.  Ru- 
dolph, S.  Winterath,  A.  Grubbs,  W.  Elliott,  B.  Martin,  and  J. 
Reddell:  Porcupine  Cave,  5  km  N  Columbia,  4  Feb.  1979,  28 
Feb.  1979,  D.  Rudolph,  S.  Winterath,  and  B.  Martin;  mine  on 
ridge,  4.5  km  N  Columbia,  22  Feb.  1979,  D.  Rudolph,  S.  Win- 
terath, and  B.  Martin;  Banksula  Cave,  6  km  N  Columbia,  21 
Mar.  1979,  D.  Rudolph,  B.  Martin,  S.  Winterath,  and  W.  El- 
liott; McLean's  Cave,  4.5  km  N  Columbia,  13  May  1967,  14 
Dec.  1977,  18  Dec.  1977,  6  Mar.  1979,  27  Mar.  1979,  2  Apr. 

1979,  K.  Horn,  T.  Briggs,  W.  Elliott,  A.  Grubbs,  S.  Winterath, 
D.  Rudolph,  and  B.  Martin;  Transplant  Mine,  3  km  N  Colum- 
bia, 17  Apr.  1979,  D.  Rudolph,  S.  Winterath,  and  E.  vanlngen. 

NOTES. — The  migration  of  Banksula  grahami 
into  two  of  the  mine  tunnels  recorded  above  is 
the  only  known  example  of  probable  interstitial 
movement  by  a  species  of  Banksula. 

Banksula  in  el  ones  Briggs 

Banksula  melones  BRIGGS  1974:8. 

RECORDS. — Calaveras  County:  Cave  of  Skulls,  5  km  NW 
Columbia,  16  Apr.  1977,  29  Mar.  1979,  T.  Briggs,  D.  Rudolph, 
B.  Martin,  S.  Winterath,  W.  Elliott,  and  J.  Reddell;  Quail 
(Gerritt's)  Cave,  5  km  NW  Columbia,  3  Jul.  1975,  17  Apr. 
1977,  30  Mar.  1979,  W.  Rauscher,  D.  Cowan,  B.  Martin,  and 
S.  Winterath;  Barren  Cave,  5  km  NW  Columbia,  30  Mar.  1979, 
D.  Rudolph,  B.  Martin,  S.  Winterath,  W.  Elliott,  and  J.  Red- 
dell; Beta  Cave,  5  km  NW  Columbia,  7  May  1977,  29  Mar. 
1979,  A.  Grubbs,  D.  Broussard,  S.  Winterath,  D.  Rudolph, 
W.  Elliott,  J.  Reddell,  and  B.  Martin;  Poison  Oak  Cave,  5  km 
NW  Columbia,  29  Mar.  1979,  D.  Rudolph,  B.  Martin,  S.  Win- 
terath, W.  Elliott,  and  J.  Reddell;  Coral  Cave,  5  km  N  Colum- 
bia, 24  Feb.  1978,  22  Mar.  1979,  A.  Grubbs,  D.  Rudolph,  B. 
Martin,  S.  Winterath,  W.  Elliott,  and  J.  Reddell;  Bryden's 
Cave,  5  km  NW  Columbia,  29  May  1977,  A.  Grubbs  and  B. 
Hopkins;  Cone  Cave,  5  km  NW  Columbia,  30  Mar.  1979,  D. 
Rudolph,  B.  Martin,  S.  Winterath,  W.  Elliott,  and  J.  Reddell; 
Eagle  View  Cave  No.  2,  5  km  NW  Columbia,  29  Mar.  1979, 
D.  Rudolph,  B.  Martin,  S.  Winterath,  W.  Elliott,  and  J.  Red- 


BRIGGS  AND  UBICK:  CAVE  HARVESTMEN 


321 


FIGURE  5.    Banksula  elliotti  Briggs  and  Ubick,  new  species.  Lateral  view  of  body  and  left  palpus  of  male  holotype. 


dell;  Lost  Piton  Cave,  6.5  km  w  Columbia,  6  May  1977,  26 
Mar.  1979,  A.  Grubbs,  N.  Boice,  D.  Broussard,  S.  Winterath, 
W.  Elliott,  and  J.  Reddell;  Bone  Cave,  6  km  w  Columbia,  20 
Apr.  1980,  T.  Briggs  and  D.  Ubick.  Tuolumne  County:  Quarry 
(McNamee's)  Cave,  3  km  NW  Columbia,  24  May  1969,  G. 
Leung,  W.  Rauscher,  and  T.  Briggs;  Gate  Pit  Cave,  3  km  NW 
Columbia,  1  Apr.  1979,  D.  Rudolph;  Mine  Cave,  3  km  NW 
Columbia,  1  Apr.  1979,  D.  Rudolph,  B.  Martin,  and  S.  Win- 
terath; Scorpion  Cave,  4.5  km  N  Columbia,  8  Feb.  1979,  25 
Mar.  1979,  D.  Rudolph,  S.  Winterath,  D.  Cowan,  and  T. 
Briggs;  Vulture  Cave,  4.5  km  N  Columbia,  10  Feb.  1979,  17 
Feb.  1979,  22  Feb.  1979,  19  Mar.  1979,  S.  Winterath,  D.  Ru- 
dolph, B.  Martin,  and  W.  Elliott;  McLean's  Cave,  4.5  km  N 
Columbia,  13  May  1967,  17  June  1967,  24  June  1967,  14  Dec. 
1977,  18  Dec.  1977,  6  Mar.  1979,  27  Mar.  1979,  K.  Horn,  T. 
Briggs,  V.  Lee,  W.  Elliott,  A.  Grubbs,  S.  Winterath,  D.  Ru- 
dolph, and  B.  Martin;  Transplant  Mine,  3  km  N  Columbia,  17 
Apr.  1979,  D.  Rudolph,  S.  Winterath,  and  E.  vanlngen. 

NOTES. — Some  of  the  specimens  collected  in 
Vulture  Cave  and  Bone  Cave  were  found  under 
rocks  in  the  twilight  zone.  All  other  collections 
were  made  in  permanently  dark  regions  of 
caves. 

ECOLOGICAL  REVIEW 

Several  ecological  studies  on  Banksula  (Briggs 
1975;  Elliott  1978;  Rudolph  1979)  have  been  con- 
ducted in  McLean's  Cave  at  the  confluence  of 
the  Middle  Fork  and  the  South  Fork  of  the  Stan- 
islaus River  (approximate  elevation,  300  m). 
This  cave  is  one  of  the  largest  in  the  Calaveras 
limestone  of  the  central  Sierra  Nevada  and  con- 
tains the  only  sympatric  populations  of  Bank- 
sula species.  Banksula  melones  and  B.  grahami 
are  mixed  in  habitats  near  the  base  of  a  broad 
talus  cone  formed  by  debris  gradually  moving 
through  two  small  entrances  to  the  lowest  levels 
of  the  cave.  Because  there  is  no  flowing  water 
in  the  cave,  the  talus  cone  and  deep-penetrating 
roots  are  important  food  sources  for  the  inhab- 
itants. The  cave  temperature  ranges  from  14  C 


to  16  C  and  the  humidity  from  82%  to  97%  (El- 
liott 1978).  Specimens  of  Banksula  melones  and 
B.  grahami  were  found  under  rocks  or  wander- 
ing on  the  floor  or  walls.  They  were  rarely  cap- 
tured in  baited  pitfall  traps  (Briggs  1975;  Elliott 
1978).  Although  biased  by  more  intensive  winter 
collecting,  records  from  McLean's  and  other 
caves  suggest  greatest  activity  in  winter  and 
spring. 

At  least  30  species  of  arthropods,  most  of 
which  were  listed  by  Elliott  (1978),  coexist  with 
Banksula  melones  and  B.  grahami  in  McLean's 
Cave.  Elliott  (1978)  identified  some  species,  but 
many  have  not  yet  been  identified.  His  feeding 
experiments  showed  that  captive  Banksula, 
kept  in  McLean's  Cave,  will  eat  live  Collembola 
but  not  equally  abundant  Psocoptera.  Rudolph 
(1979)  maintained  eight  immature  Banksula  in 
McLean's  Cave  using  Collembola  as  food;  three 
molted  within  43  days.  One  adult  and  two  im- 
mature individuals  maintained  at  the  same  time 
without  food  survived. 

Rudolph  (1979)  searched  the  mine  tunnel  into 
which  Briggs  and  Elliott  transplanted  Banksula 
melones,  B.  grahami,  and  miscellaneous  arthro- 
pods from  McLean's  Cave.  The  transplanted 
population,  which  included  a  few  hundred 
Banksula,  appeared  to  be  reproducing.  The  fu- 
ture of  this  transplant  may  depend  on  how  well 
conditions  in  the  mine  duplicate  conditions  in 
McLean's  Cave.  If  they  both  flourish,  the  trans- 
planted Banksula  species  may  demonstrate  that 
their  sympatry  in  a  small  cave  is  stable. 

BlOGEOGRAPHY 

The  distribution  of  Banksula  species  is  similar 
to  the  invertebrate  troglobite  distributions  re- 
ported from  caves  in  the  Appalachian  Mountains 


322 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  1 1 


of  the  eastern  United  States  in  that  these  cave 
species  tend  to  be  bounded  within  "karst  is- 
lands," within  which  subterranean  dispersal  and 
genetic  communication  readily  occurs  (Barr 
1967;  Culver  et  al.  1973).  Such  karst-island  spe- 
ciation  exists  for  Banksula  in  the  Sierra  Nevada 
if  the  limestone  outcrops  are  widely  separated. 
Thus,  the  species  B.  galilei  Briggs,  B.  calif or- 
nica  (Banks),  B.  rudolphi  new  species,  B. 
grubbsi  new  species,  and  B.  tuolumne  Briggs 
occupy  karst  that  is  separated  by  many  kilo- 
meters of  nonporous  rock  (Fig.  1).  The  isolation 
of  these  cave  species  is,  therefore,  more  com- 
plete than  that  of  the  species  in  the  Appalachian 
caves.  The  area  of  the  karst  in  which  B.  galilei, 
B.  californica,  and  B.  rudolphi  are  found  is 
0.02-0.1  km2  and  is  considerably  smaller  than 
the  smallest  karst  island  (10  km2)  studied  in  the 
eastern  United  States  (Culver  et  al.  1973).  Fur- 
thermore, quarrying  shows  that  these  small  out- 
crops have  small  volume  and  little  internal  water 
flow. 

All  known  Banksula  species  are  allopatric  ex- 
cept for  B.  melones  and  B.  grahami  in  Mc- 
Lean's Cave.  This  suggests  that  sympatric 
species  compete,  and  the  sympatry  in  McLean's 
Cave  is  a  result  of  a  recent  invasion.  About  half 
of  the  70  or  so  Calaveras  Formation  caves  care- 
fully checked  yielded  a  species  of  Banksula. 
Thirty-one  of  these  are  located  in  the  karst  of 
the  Stanislaus  River.  Eighteen  apparently  hab- 
itable caves  in  this  region  did  not  contain  Bank- 
sula (Rudolph  1979).  The  absence  of  harvest- 
ment  in  caves  located  in  karst  outside  of  the 
Stanislaus  River  region  may  be  due  to  an  uneven 
distribution  of  their  epigean  ancestors. 

The  distribution  of  the  four  species  of  Bank- 
sula (B.  melones,  B.  grahami,  B.  martinorum, 
and  B.  elliotti}  that  occupy  the  karst  of  the  Stanis- 
laus region  is  not  readily  explained.  Volcanic 
rock  divides  Coyote  Creek  (west  of  the  Stanis- 
laus River)  from  the  Stanislaus  River.  It  forms 
an  irregular  barrier  between  B.  elliotti  and  B. 
melones,  and  separates  B.  melones  from  B.  gra- 
hami southeast  of  the  Stanislaus  River.  Several 
continuous  bands  of  amphibolite  divide  the  re- 
maining limestone  regions  into  parallel  lenses  in 
which  allopatric  populations  of  B.  melones  and 


B.  grahami  occupy  most  of  the  available  cav- 
erns. These  populations  seem  to  split  into  east- 
ern and  western  clusters  that  do  not  correlate 
well  with  geology  or  water  systems.  The  karst 
that  contains  the  easternmost  population  of  B. 
elliotti  is  isolated  by  one  kilometer  of  metavol- 
canic  rock  from  the  karst  that  contains  the  rest 
of  B.  elliotti  and  B.  martinorum.  Coyote  Creek, 
the  much  larger  Stanislaus  River,  and  the  south 
fork  of  the  Stanislaus  that  branches  to  the  east 
have  no  apparent  effect  on  the  distribution  of 
species  of  Banksula.  The  four  Stanislaus  species 
show  increasing  cavernicolous  specialization  in 
relative  eye  loss,  from  B.  melones  with  the  larg- 
est eyes,  to  B.  elliotti  with  missing  retinae,  and 
B.  martinorum  with  complete  eye  loss.  One 
might  infer  that  adaptive  radiation  began  with  a 
B.  melones-\ike  ancestor  and  progressed  to  B. 
elliotti.  If  correct,  one  must  explain  how  the  in- 
creasingly troglobitic  species  B.  grahami  and  B. 
elliotti  were  able  to  cross  geologic  and  drainage 
barriers.  Further  studies  on  the  harvestmen  of 
the  Stanislaus  River  region  are  needed  to  explain 
this  unexpected  distribution  of  species.  It  is  re- 
grettable that  this  portion  of  the  river  will  soon 
be  inundated  by  the  New  Melones  Reservoir  and 
some  of  the  harvestmen  caves  will  be  lost. 

LITERATURE  CITED 

BARR,  T.  C.,  JR.   1967.  Observations  on  the  ecology  of  caves. 

Am.  Nat.  101(922):475^t91. 
BRIGGS,  T.  S.    1968.  Phalangids  of  the  laniatorid  genus  Sital- 

cina  (Phalangodidae:Opiliones).   Proc.   Calif.   Acad.   Sci. 

36(0:3-7. 
.    1971.  Relict  harvestmen  from  the  Pacific  Northwest. 

Pan-Pac.  Entomol.  47(3):  168-169. 
.   1974.  Phalangodidae  from  caves  in  the  Sierra  Nevada 


(California)  with  a  redescription  of  the  type  genus.  Occas. 
Pap.  Calif.  Acad.  Sci.  108:1-15. 
.    1975.    Biological  transplant  project,  New  Melones 


Lake,  California— final  report.  Serial  #DACW05-75-P-1845, 
U.S.  Army  Corps  of  Eng.,  Sacramento  Dist.,  Calif. 

CULVER,  D.,  J.  R.  HOLSINGER,  AND  R.  BAROODY.  1973.  To- 
ward a  predictive  cave  biogeography:  the  Greenbrier  Valley 
as  a  case  study.  Evolution  27(4): 689-695. 

ELLIOTT,  W.  R.  1978.  Final  report  on  the  New  Melones  cave 
harvestman  transplant.  Contract  #DACW05-78-C-0007, 
U.S.  Army  Corps  of  Eng.,  Sacramento  Dist.,  Calif. 

RUDOLPH,  D.  C.  1979.  Final  report  on  the  status  of  the  Me- 
lones cave  harvestman  in  the  Stanislaus  River  drainage. 
Contract  #14-16-0009-79-009,  U.S.  Fish  Wildl.  Serv., 
Wash.  D.C. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  12,  pp.  323-339,  34  figs.  June  24,  1981 


TROPICAL  EASTERN  PACIFIC  LIMPETS  OF  THE  FAMILY 

ACMAEIDAE  (MOLLUSCA,  ARCHAEOGASTROPODA): 

GENERIC  CRITERIA  AND  DESCRIPTIONS  OF  SIX 

NEW  SPECIES  FROM  THE  MAINLAND  AND 

THE  GALAPAGOS  ISLANDS 

By 
David  R.  Lindberg* 

Center  for  Coastal  Marine  Studies,  University  of  California, 
Santa  Cruz,  California  95064 

and 
James  H.  McLean 

Section  of  Malacology,  Natural  History  Museum  of  Los  Angeles  County, 
Los  Angeles,  California  90007 


ABSTRACT:  We  define  genera  on  conservative  shell  structure  characters  and  on  qualitative  radular  charac- 
ters. Lottia,  previously  considered  monotypic,  is  expanded  to  include  Panamic  species  with  a  secondary  gill 
(branchial  cordon)  formerly  assigned  to  Scurria.  Scurria  has  a  similar  gill,  but  the  shell  structure  differs.  The 
new  species  Notoacmea  ubiquita  from  Mexico  and  N.  pumila  from  Ecuador  are  small-shelled  allopatric  species 
with  radular  teeth  modified  for  feeding  on  coralline  algae.  Two  new  species  of  Notoacmea  (N.  rothi  and  N. 
immaculata),  endemic  to  the  Galapagos  Islands,  constitute  a  species  pair  differing  chiefly  in  radular  features: 
the  radular  teeth  of  \ .  immaculata  are  adapted  for  feeding  on  calcareous  algae;  those  of  V.  rothi  for  noncal- 
careous  algae.  A  pair  of  endemic  new  species  of  Lottia  from  the  Galapagos  Islands  (/,.  mimica  and  /..  smithi) 
also  differ  mainly  in  radular  characters,  l.ottiu  mimica  is  a  noncalcareous-alga  feeder  and  /,.  smithi  is  a  calcar- 
eous-alga feeder.  These  four  endemic  species  are  the  principal  acmaeid  limpets  of  the  Galapagos.  Two  mainland 
species,  Notoacmea  filosa  and  Lottia  mesoleuca,  are  known  only  sporadically  from  the  Galapagos  Islands. 

INTRODUCTION 

-,,     ,  •        MI  c     described  herein — Notoacmea  ubiquita   and 

The  last  comprehensive,  illustrated  review  of 

the  Acmaeidae  of  the  tropical  eastern  Pacific      Lottm  ^mica-were  recognized  as  new  but 
was  given  by  McLean  (1971).  At  the  time  of     *ere  not  Bribed.  The  «fnenc  placement  of 

preparation  of  that  account,  two  of  the  species      these  tw°  sPec,ies  was  puzz'!n8  because,  *<*  *ad 

previously  unknown  combinations  of  radular, 

shell,  and  gill  characters. 

,  ,T  Further  study  of  generic  relationships  has  now 

*  Research  Associate,  Department  of  In  vertebrate  Zoology,  J        ° 

California  Academy  of  Sciences,  San  Francisco,  California       provided  a  basis  for  the  convincing  allocation  of 

94118.  these  species.  Although  a  full  review  of  generic 

[323] 


324 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  12 


criteria  in  the  family  is  beyond  the  scope  of  this 
paper,  we  include  some  discussion  pertaining  to 
the  Panamic  species.  A  major  distinction  has 
become  apparent  between  Lottia  Sowerby, 
1834,  and  Scurria  Gray,  1847,  two  genera  having 
a  secondary  gill  (accessory  gill  lappets  on  the 
mantle  margin).  These  two  genera  are  redefined 
here,  the  name  Lottia  thus  being  made  available 
for  use  for  some  tropical  species  previously  con- 
sidered to  belong  to  Scurria. 

A  closer  examination  of  the  acmaeids  of  the 
Galapagos  Islands  has  resulted  in  the  recogni- 
tion of  four  new  endemic  species,  representing 
two  species  pairs  wherein  the  principal  differ- 
ences are  in  radular  tooth  morphology.  The  shell 
characters  of  each  pair  are  insufficiently  distinct 
to  permit  reliable  identification  by  shell  alone. 
Radular  characters  in  the  Acmaeidae  have  been 
found  by  all  workers  to  be  species-specific.  In 
no  species  has  ontogenetic  or  situs  variation  in 
radulae  been  found.  Similar  shell  morphologies 
have  been  reported,  however,  in  both  conge- 
neric and  noncongeneric  species  of  Acmaeidae 
(McLean  1966;  Lindberg  1979).  We  therefore 
consider  each  radular  morphotype  to  represent 
a  separate  species.  Because  the  shell  characters 
of  each  pair  are  insufficiently  distinct  to  permit 
reliable  identification,  both  species  are  dis- 
cussed in  a  combined  discussion  section  follow- 
ing their  formal  descriptions. 

Abbreviations  are  as  follows:  AHF,  Allan 
Hancock  Foundation,  University  of  Southern 
California,  Los  Angeles  (collection  on  loan  to 
LACM);  AMNH,  Department  of  Invertebrates, 
American  Museum  of  Natural  History,  New 
York;  ANSP,  Department  of  Malacology,  Acad- 
emy of  Natural  Sciences,  Philadelphia;  CAS, 
Department  of  Invertebrate  Zoology,  California 
Academy  of  Sciences,  San  Francisco;  LACM, 
Section  of  Malacology,  Natural  History  Mu- 
seum of  Los  Angeles  County,  Los  Angeles; 
MCZ,  Museum  of  Comparative  Zoology,  Har- 
vard University,  Cambridge;  SU,  Stanford  Uni- 
versity, Stanford  (collection  on  loan  to  CAS); 
USNM,  Division  of  Mollusks,  U.S.  National 
Museum  of  Natural  History,  Washington,  D.C. 

GENERIC  CRITERIA  FOR  THE 
PANAMIC  ACMAEIDAE 

Generic  assignments  in  McLean's  (1971)  re- 
view were  based  on  shell  sculpture,  presence  or 


absence  of  the  secondary  gill,  and  whether  mar- 
ginal radular  teeth  are  represented  by  two  pairs 
of  fully  developed  teeth,  a  single  pair  of  rudi- 
mentary teeth  (uncini),  or  are  absent  altogether. 
We  now  realize  that  a  system  based  on  these 
three  characters  alone  is  not  adequate. 

Christiaens  (1975)  proposed  a  generic  classi- 
fication of  the  Acmaeidae  in  which  tooth  shape 
and  configuration  were  especially  emphasized. 
He  felt  some  genera  had  three  pairs  of  lateral 
teeth  and  some  two,  the  latter  group  having  a 
bicuspid  second  lateral  tooth.  We  maintain  that 
all  acmaeids  have  three  pairs  of  lateral  teeth.  We 
fail  to  see  how  the  third  lateral  tooth  can  be 
interpreted  as  part  of  the  second,  because  in  all 
acmaeid  radulae  we  have  examined,  we  find  that 
the  ventral  plates  of  the  radular  ribbon  have 
three  lateral  plate  components,  one  correspond- 
ing to  each  lateral  tooth.  We  believe  that  the 
reduction  of  the  outermost  tooth  that  occurs  in 
some  species  is  a  result  of  dietary  specialization. 
Relation  of  diet  to  tooth  shape  was  discussed  by 
McLean  (1966),  and  we  are  now  aware  of  similar 
tooth  shape  and  configuration  in  species  of  di- 
verse genera.  We  do  not  consider  lateral  tooth 
shape  to  be  useful  as  a  generic  character. 

We  continue  to  maintain  full  generic  separa- 
tion of  species  groups  in  which  the  marginal 
teeth  have  three  possible  expressions:  (1)  two 
pairs  of  fully  functional  marginals  (Patelloida 
Quoy  and  Gaimard,  1834);  (2)  a  single  pair  of 
marginal  remnants  or  uncini  (Collisella  Dall, 
1871;  Lottia  Sowerby,  1834;  Scurria  Gray, 
1847);  and  (3)  no  marginals  or  uncini  (Acmaea 
Eschscholtz,  1833;  Notoacmea  Iredale,  1915; 
Problacmaea  Golikov  and  Kussakin,  1972; 
Rhodopetala  Dall,  1921;  and  Tectura  Gray, 
1847).  We  therefore  disagree  with  Christiaens' s 
ranking  of  Notoacmea  as  a  subgenus  of  Colli- 
sella. 

Recent  work  by  Lindberg  (1976,  1978)  has  em- 
ployed shell  structure  characters  first  used  for 
patellacean  limpets  by  MacClintock  (1967).  We 
believe  that  the  relationships  suggested  by  shell 
structure  are  conservative  and  are  basic  to  a 
modern  classification  of  the  family. 

We  are  now  inclined  to  define  genera  using 
shell  structure,  branchial  characters,  radula  bas- 
al plate  structure,  and  the  three  possibilities  for 
marginal  teeth  listed  above,  recognizing  that 
shell  sculpture  characters  are  convergent  in  all 
genera  and  that  lateral  tooth  shape  is  likewise 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


325 


FIGURES  1—4.  Ventral  views  of  preserved  specimens  of  Lottia  and  Scurria  species  showing  secondary  gill  in  relation  to 
head.  FIGURE  1.  Lottia  gigantea,  Isla  de  Guadalupe,  Mexico  (LACM  55618).  FIGURE  2.  Lottia  mesoleuca,  Bahia  Tenacatita, 
Jalisco,  Mexico  (LACM  66-55).  FIGURE  3.  Lottia  mimica  new  species,  paratype,  Academy  Bay,  Isla  Santa  Cruz,  Galapagos 
Islands,  Ecuador  (LACM  1926).  FIGURE  4.  Scurria  scurra,  Punta  El  Lacho,  Santiago  Province,  Chile  (LACM  75-32). 


convergent  and  widely  variable  interspecifically 
(although  not  intraspecifically).1 

Acmaea  and  Notoacmea  differ  in  lateral  plate 
morphology.  In  Acmaea  the  lateral  plates  are 
similar  in  size  and  shape  and  are  arranged  in  a 
posteriorly  diverging  V-configuration.  In  No- 
toacmea the  lateral  plates  are  unequal  in  size 
and  shape — the  first  and  second  lateral  plates 
tend  to  lie  in  the  same  line  and  the  third  lateral 
plates  are  always  lateral  and  slightly  posterior 
to  the  second  lateral  plates. 


1  In  his  discussion  of  Notoacmea  fascicularis  (Menke, 
1851)  McLean  (1971:327)  alluded  to  two  different  radular 
types  within  that  species,  suggesting  that  a  "complex  involv- 
ing more  than  one  species"  was  a  possibility.  Lindberg  will 
report  separately  on  the  two  species  of  the  N.  fascicularis 
complex. 


Acmaea  and  Tectura  also  differ  in  lateral  plate 
morphology.  In  both  genera  dentition  consists 
of  three  pairs  of  equal-sized  and  equal-shaped 
lateral  teeth;  however,  in  Tectura  the  lateral 
plates  that  support  these  teeth  are  complex  and 
similar  in  shape  and  position  to  those  found  in 
the  genus  Collisella. 

All  species  of  Acmaea  are  known  to  feed  on 
coralline  algae  and  have  blunt,  equal-sized  teeth. 
Some  of  the  tropical  eastern  Pacific  and  Carib- 
bean species  of  Notoacmea  are  now  known  to 
have  teeth  similarly  blunt  and  of  equal  size. 
These  species  may  also  be  coralline  alga  feeders. 
Three  of  the  four  new  species  of  Notoacmea 
described  in  this  paper  (N.  ubiquita,  N.  pumila, 
and  N.  immaculatd)  have  blunt  equal-sized  lat- 
eral teeth.  The  other  new  species  of  Notoac- 
mea, N.  rothi,  has  the  outermost  lateral  tooth 


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greatly  reduced  and  the  first  two  pairs  more 
elongate,  which  is  the  pattern  characteristic  of 
most  temperate  and  tropical  species  of  Notoac- 
mea. 

Similar  modification  of  the  lateral  teeth  for 
feeding  on  coralline  algae  is  known  in  some  trop- 
ical species  of  Collisella.  Eastern  Pacific  species 
of  this  uncinate  genus  with  lateral  teeth  so  mod- 
ified are:  C.  atrata  (Carpenter,  1857),  C.  discors 
(Philippi,  1849),  C.  mitella  (Menke,  1847),  and 
C.  pediculus  (Philippi,  1846). 

Two  generic  names  have  been  used  for  ac- 
maeid  limpets  in  which  there  is  a  secondary  gill 
(branchial  cordon)  in  addition  to  the  normal  ac- 
maeid  ctenidium:  Lottia  Sowerby,  1834  (type- 
species  L.  gigantea  Sowerby,  1834),  and  Scur- 
ria  Gray,  1847  (type-species  Patella  scurra 
Lesson,  1830).  Lottia  has  usually  been  consid- 
ered monotypic,  with  the  single  Californian 
species  L.  gigantea.  It  has  been  diagnosed  (Dall 
1871)  as  having  a  secondary  gill  incomplete  or 
interrupted  in  front  of  the  head,  whereas  in 
Scurria  the  gill  is  complete  or  continuous.  The 
radular  dentition  in  both  genera  consists  of  three 
pairs  of  lateral  teeth  and  one  pair  of  uncini. 

We  have  examined  the  secondary  gill  in  living 
and  preserved  specimens  of  L.  gigantea  and 
find  that  many  specimens  have  a  greatly  reduced 
but  distinct  gill  in  front  of  the  head  (Fig.  1).  The 
secondary  gill  of  a  tropical  eastern  Pacific 
species  usually  assigned  to  Scurria,  S.  mesoleu- 
ca  (Menke,  1851),  is  normally  much  less  prom- 
inent in  front  of  the  head  than  along  the  sides 
(Fig.  2).  The  secondary  gill  of  Lottia  mimica, 
new  species  (Fig.  3),  is  also  much  reduced  in 
front  of  the  head.  The  secondary  gill  of  Scurria 
scurra  (Fig.  4)  is  complete  over  the  head,  but  it 
is  also  somewhat  reduced  in  prominence  in  this 
region.  We  therefore  do  not  regard  the  reduction 
of  the  secondary  gill  near  the  head  as  a  useful 
generic  character. 

MacClintock  (1967)  found  that  Scurria  in  the 
Peruvian  faunal  province  differ  in  shell  structure 
from  other  eastern  Pacific  species  with  the  sec- 
ondary gill.  The  Peruvian  Scurria  species  are  in 
MacClintock's  shell  structure  "group  3," 
whereas  Lottia  gigantea  and  the  two  species 
placed  by  McLean  (1971)  in  Scurria  (S.  meso- 
leuca  and  5.  stipulata  (Reeve,  1855))  are  in  shell 
structure  "group  1"  (along  with  most  other 
species  of  Collisella  and  Notoacmea).  Because 
we  believe  that  shell  structure  is  more  conser- 


vative than  branchial  characters,  and  we  place 
even  less  emphasis  on  shell  sculpture  and  col- 
oration, we  infer  that  the  Panamic  acmaeids  pre- 
viously assigned  to  Scurria  are  more  closely  re- 
lated to  L.  gigantea  than  to  Scurria.  The  two 
Panamic  species  plus  L.  mimica  and  L.  smithi 
described  in  this  paper  are  therefore  assigned  to 
Lottia.  Lottia  is  redefined  to  include  uncinate 
species  in  shell  structure  "group  1,"  with  a  sec- 
ondary gill  that  is  usually  reduced  but  not  nec- 
essarily absent  over  the  head. 

NEW  SPECIES  OF  ACMAEIDAE  FROM  THE 
TROPICAL  EASTERN  PACIFIC 

Notoacmea  ubiquita  new  species 

(Figures  5-7,  23,  29) 

There  are  two  situs  forms  of  this  species,  a 
laterally  compressed  form  and  an  oval  form.  A 
description  for  each  follows. 

Shell  (oval  form)  (Figs.  5,6):  Relatively  small 
(maximum  length  12  mm),  profile  of  medium 
height;  apex  anterior  to  center;  all  slopes  con- 
vex; large  shells  frequently  with  a  flattened  area 
posterior  to  apex;  sides  of  shell  somewhat  par- 
allel. Sculpture  of  rounded  radial  ribs,  with 
weaker  secondary  ribs  beginning  below  apex; 
ribs  extending  slightly  beyond  the  shell  edge, 
crenulating  the  aperture;  concentric  sculpture  of 
well-defined  but  nearly  microscopic,  sharply 
raised  ridges.  Exterior  translucent  white  with 
red-brown  markings  on  early  shell,  the  markings 
becoming  darker,  reticulate,  and  limited  to  rib- 
interspaces  with  growth;  ribs  white,  overlain 
with  dark  brown  radial  markings  that  may  be 
concentrated  into  lateral  rays.  Interior  margin 
white  with  dark  markings  that  correspond  to  ex- 
terior interspaces;  intermediate  area  white;  cen- 
tral area  with  yellow  stain,  exterior  markings 
visible  through  shell. 

Shell  (compressed  form)  (Fig.  7):  Lateral  pro- 
file high,  ends  raised  relative  to  sides;  all  slopes 
convex;  some  specimens  compressed  in  early 
stage,  changing  abruptly  to  oval  form.  Sculpture 
and  coloration  as  in  oval  form. 

Radula  (Figs.  23,  29):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
medial  edges  convex,  lateral  edges  straight  to 
slightly  concave,  cusps  rounded,  blunt;  second 
pair  of  lateral  teeth  posterior  and  lateral  to  first 
pair,  medial  and  lateral  edges  convex,  cusps 
rounded,  blunt;  third  lateral  teeth  lateral  to  sec- 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


327 


10 


FIGURES  5-10.  FIGURE  5.  Notoacmea  ubiquita  new  species.  Holotype,  LACM  1917.  Santiago  Peninsula,  Colima,  Mexico. 
Length  11.7  mm.  FIGURE  6.  Notoacmea  ubiquita.  LACM  54773.  Guaymas,  Sonora,  Mexico.  Length  6.9  mm.  FIGURE  7. 
Notoacmea  ubiquita.  LACM  54773.  Guaymas,  Sonora,  Mexico.  Length  5.4  mm.  FIGURE  8.  Notoacmea  pumila  new  species. 
Holotype,  LACM  1919.  Punta  Ancon,  Ecuador.  Length  4.7  mm.  FIGURE  9.  Notoacmea  pumila.  Paratype,  LACM  1920.  Punta 
Ancon,  Ecuador.  Length  3.6  mm.  FIGURE  10.  Notoacmea  pumila.  LACM  72-17.  Bahia  Jobo,  Costa  Rica.  Length  5.6  mm. 


ond  pair,  medial  edges  concave,  lateral  edges 
straight,  extending  to  edges  of  ventral  plates, 
cusps  blunt.  Marginal  teeth  lacking.  First  lateral 
plates  overlap  anterior  ribbon  segment,  postero- 
lateral  edges  concave;  second  lateral  plates  ir- 
regular, posterior  edges  convex;  third  lateral 
plates  lobate  with  lateral  lobes  extending  to 
edges  of  ventral  plates;  second  and  third  lateral 
plates  separated  by  a  partial  suture.  Ventral 


plates  closely  set  with  both  anterior  and  poste- 
rior processes. 

Animal:  Pigmentation  lacking,  snout  with  oral 
lappets. 

HOLOTYPE  DIMENSIONS. — Length  11.7,  width 
8.1,  height  3.6  mm. 

TYPE-LOCALITY. — Mexico:  Colima;  Manza- 
nillo,  Santiago  Peninsula,  Playa  Las  Hadas 
(19°05'57"N,  103°19'36"W)  (LACM  63-10),  inter- 


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tidal  zone  to  5  m.  Leg.  J.  H.  McLean  and  C. 
Tenney,  21-24  Mar.  1963,  3  specimens. 

TYPE-MATERIAL. — Holotype  (oval  form), 
LACM  1917;  2  paratypes,  LACM  1918  (both 
oval  and  compressed  forms).  The  holotype  is  the 
largest  specimen  examined. 

DISTRIBUTION. — Baja  California  Sur,  Mexico, 
from  Punta  Pequena  (26°14'N)  (LACM  71-6)  to 
Bahia  Magdalena  and  Cabo  San  Lucas,  north  in 
the  Gulf  of  California  to  Puertecitos  (30°25'N) 
(LACM  65-34)  and  Guaymas,  south  to  Bahia 
Tangola  Tangola,  Oaxaca  (15°45'N)  (AHF  215). 

MATERIAL  EXAMINED. — 58  lots,  approxi- 
mately 350  specimens,  3  radula  preparations. 

ETYMOLOGY. — The  name  is  based  on  the  Lat- 
in adverb  ubique  (everywhere).  The  species  is 
indeed  ubiquitous — shells,  at  least,  have  been 
recovered  from  sediment  residues  taken  by 
divers  from  nearly  all  LACM  station  localities 
throughout  the  range. 

DISCUSSION. — Although  Notoacmea  ubiquita 
has  been  known  for  many  years,  it  was  not  de- 
scribed earlier  because  of  uncertainties  about  its 
generic  position.  Its  shell  shape  and  sculpture 
suggested  Collisella,  and  its  lack  of  uncini  and 
subtidal  habitat  suggested  Acmaea.  We  are  now 
satisfied  to  place  it  in  Notoacmea  because  of 
the  basal  plate  configuration.  Although  most 
species  of  Notoacmea  are  finely  ribbed,  the 
rather  more  prominent  ribbing  of  N.  ubiquita  is 
not  as  strong  as  occurs  in  many  species  of  Col- 
lisella. 

Notoacmea  ubiquita  is  the  only  Panamic  ac- 
maeid  with  an  oval  form  and  a  laterally  com- 
pressed form.  Notoacmea  ubiquita  has  broader, 
stronger  ribs  than  any  other  tropical  species  of 
Notoacmea.  On  the  basis  of  shell  characters,  it 
is  more  similar  to  some  of  the  Panamic  Colli- 
sella, which  differ  in  having  an  uncinate  radula. 
The  fine  brown  concentric  markings  of  the  early 
stages  do  not  occur  in  any  other  species.  Col- 
lisella turveri  (Hertlein  and  Strong,  1951)  differs 
in  having  broader,  more  projecting  ribs.  Colli- 
sella acutapex  (Berry,  1960)  has  a  higher  shell 
profile  with  sharper,  more  prominent  ribbing, 
and  its  pattern  of  brown  lines  is  more  coalescing. 
Collisella  mitella  (Menke,  1847)  also  has  white 
ribs,  but  its  ribs  are  more  numerous  and  the  in- 
terspaces are  dark  colored.  Patelloida  semiru- 
bida  (Dall,  1914)  has  sharper  radial  and  concen- 
tric sculpture,  with  red  rather  than  brown 
markings;  its  radula  is  also  markedly  different, 


having  two  pairs  of  marginal  teeth  per  ribbon 
segment. 

Large  lots  show  a  complete  series  of  possible 
shell  shapes  between  the  elevated  narrow  forms 
with  raised  ends  and  the  low  oval  forms.  Some 
shells  have  an  early  compressed  phase,  with  lat- 
er growth  stages  like  the  oval  form;  some  rela- 
tively large  shells  are  angulate  at  the  sides,  giv- 
ing the  shell  a  flat-topped  appearance.  Color 
variation  is  relatively  minor;  one  color  variant 
characteristic  of  specimens  from  Jalisco  is  or- 
dinary in  early  stages,  changing  to  solid  maroon 
at  later  stages.  Largest  shells  seen  are  from  Ja- 
lisco and  Colima;  specimens  from  localities  in 
the  Gulf  of  California  attain  about  two-thirds  the 
size  of  southern  specimens. 

Notoacmea  ubiquita  has  features  in  common 
with  two  more  northern  species,  Collisella 
triangularis  (Carpenter,  1864)  and  Tectura  ro- 
sacea  (Carpenter,  1864),  both  of  which  differ  in 
lacking  the  radial  ribbing.  All  three  species  have 
laterally  compressed  forms,  are  primarily  sub- 
tidal,  and  have  equal-sized  lateral  teeth  adapted 
for  feeding  on  calcareous  algae.  In  C.  triangu- 
laris the  compressed  form  predominates,  where- 
as in  T.  rosacea  the  oval  form  is  more  abundant, 
but  in  both  species  the  compressed  forms  occur 
on  branching  coralline  algae  and  the  oval  forms 
occur  on  crustose  coralline  algae,  and  all  inter- 
mediate conditions  are  known.  Although  we 
have  not  directly  observed  the  compressed  form 
of  N.  ubiquita  on  branching  coralline  algae,  it 
probably  so  occurs,  judging  from  its  ability  to 
change  from  compressed  to  oval  during  growth, 
which  implies  a  change  of  situs. 

Notoacmea  pumila  new  species 

(Figures  8-10,  24,  30) 

Shell  (Figs.  8-10):  Small  (maximum  length  7 
mm);  profile  medium-high;  apex  anterior  to  cen- 
ter; anterior  slope  straight  to  convex,  lateral 
slopes  convex;  usually  encrusted  with  coralline 
algae.  Sculpture  of  fine,  sharp  radial  ribs  origi- 
nating below  apex,  secondary  ribs  arising  in  the 
interspaces,  not  reaching  thickness  of  primary 
ribs.  Aperture  oval,  not  crenulate.  Color  pattern 
independent  of  ribbing:  most  frequently  white 
near  apex,  gray  at  margin,  with  6  to  10  white 
rays  in  a  stellate  pattern,  some  rays  not  reaching 
margin;  some  specimens  with  fine  brown  lines 
bordering  white  rays  and  fine  brown  lines  that 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


329 


produce  a  concentric  network.  Interior  translu- 
cent white,  showing  the  exterior  pattern. 

Radula  (Figs.  24,  30):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
medial  edges  convex,  lateral  edges  concave, 
cusps  rounded.  Second  pair  of  lateral  teeth  pos- 
terior to  first  pair,  medial  edges  convex,  lateral 
edges  straight,  cusps  rounded.  Third  pair  of  lat- 
eral teeth  positioned  posterior  and  lateral  to  sec- 
ond pair,  medial  edges  convex,  lateral  edges 
straight  to  slightly  concave.  Third  laterals 
broader  than  second  laterals,  with  lateral  exten- 
sions to  edges  of  ventral  plates;  cusps  rounded. 
Marginal  teeth  lacking.  First  lateral  plates  irreg- 
ular, anterior  portions  overlapping  anterior  rib- 
bon segments;  second  lateral  plates  elongate, 
ovoid;  third  lateral  plates  triangular,  with  con- 
vex posterior  edge.  Ventral  plates  with  strong 
anterior  and  posterior  processes.  Lateral  por- 
tions with  strong  sutures  parallel  to  edges. 

Animal:  Pigmentation  lacking,  oral  fringe  sim- 
ple. 

HOLOTYPE  DIMENSIONS. — Length  4.7,  width 
3.3,  height  2.0  mm. 

TYPE-LOCALITY. — Ecuador:  Santa  Elena 
Peninsula;  Punta  Ancon,  north  and  south  sides 
(2°20'S,  80°54'W),  intertidal  zone.  Leg.  J.  H. 
McLean  and  D.  Shasky,  6-7  Mar.  1970  (LACM 
70-11,  70-12),  72  LACM  specimens,  12  Shasky 
specimens. 

TYPE-MATERIAL.— Holotype,  LACM  1919, 
paratypes,  LACM  1920;  paratypes  have  also 
been  deposited  in  the  collections  of  CAS  and 
USNM,  and  in  Shasky  collection  (Redlands, 
California). 

DISTRIBUTION. — El  Velero,  Nicaragua 
(12°01'N)  (LACM  74-86),  south  to  Ecuador 
(type-locality).  There  are  numerous  dead  spec- 
imens from  Bahia  Salinas,  Costa  Rica  (11°02'N, 
85°45'W)  (LACM  72-17,  72-19);  two  specimens 
only  from  Panama  at  San  Carlos  (8°29'N, 
79°57'W)  (LACM  75-55),  and  a  number  of  lo- 
calities in  Ecuador  collected  by  D.  Shasky. 

MATERIAL  EXAMINED. — 19  lots,  approxi- 
mately 200  specimens,  3  radula  preparations. 

ETYMOLOGY. — The  name  is  a  Latin  adjective, 
pumilus,  meaning  small  or  dwarfish — fitting  for 
this,  the  smallest  tropical  eastern  Pacific  mem- 
ber of  the  family. 

DISCUSSION. — Notoacmea  pumila  could  be 
confused  only  with  two  other  relatively  small 
forms,  N.  ubiquita  new  species  and  Patelloida 


semirubida.  It  differs  from  the  first  in  having 
much  sharper  ribbing  and  not  being  compressed. 
Although  both  N.  pumila  and  P.  semirubida 
have  fine  sharp  radial  ribs,  N.  pumila  lacks  the 
sharp  concentric  sculpture  and  pink  markings  of 
P.  semirubida. 

The  radula  of  N.  pumila  is  similar  to  that  of 
two  new  species  described  herein,  N.  ubiquita 
and  N.  immaculata.  It  differs  from  both  by  hav- 
ing a  complete  rather  than  partial  suture  be- 
tween the  second  and  third  lateral  plates  and 
having  strong  ventral  plate  sutures  parallel  to 
the  lateral  edges.  The  ventral  plates  of  N.  pum- 
ila have  anterior  and  posterior  processes  which 
N.  immaculata  lacks,  and  the  third  lateral  plates 
are  triangular  rather  than  biformed  as  in  N.  im- 
maculata and  N.  ubiquita.  The  radula  of  N. 
pumila  differs  from  that  of  P.  semirubida  by 
lacking  marginal  teeth. 

Large  lots  show  similar  color  patterns  both  in 
the  material  from  Costa  Rica  and  from  stations 
in  Ecuador.  A  small  percentage  of  specimens 
change  with  growth  from  dark  rayed  to  solid 
dark  (see  Fig.  9);  fewer  specimens  are  rayed 
only  with  brown  linear  markings  and  fine  brown 
reticulate  markings.  Shell  proportions  vary  only 
slightly. 

Notoacmea  pumila  undoubtedly  feeds  on  cor- 
alline algae — the  lateral  teeth  are  blunt  and  of 
equal  size.  Living  specimens  have  been  collect- 
ed in  the  low  intertidal  zone  and  the  species 
probably  also  occurs  in  the  immediate  subtidal 
zone  on  coralline-encrusted  rocks. 

Notoacmea  rothi  new  species 

(Figures  11-13,  25,  31) 

Shell  (Figs.  11-13):  Size  medium  (maximum 
length  20  mm),  height  medium;  apex  anterior  to 
center;  all  slopes  convex,  aperture  ovoid.  Sculp- 
ture of  unequal  riblets  and  concentric  growth 
lines;  one  to  three  secondary  riblets  between 
each  two  primary  riblets.  Exterior  dark  gray 
with  scattered  white  markings;  apical  pattern 
tessellate;  white  markings  often  aligned  in  lateral 
rays  that  define  a  broad,  dark  posterior  ray.  In- 
terior margin  broad,  dark,  streaked  with  white 
corresponding  to  exterior  pattern;  intermediate 
area  blue- white;  central  area  blue- white  with 
brown  stain. 

Radula  (Figs.  25,  31):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
medial  edges  convex,  lateral  edges  slightly  con- 


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16 


FIGURES  11-16.  Notoacmea  from  the  Galapagos  Islands.  FIGURE  11.  Notoacmea  rothi  new  species.  Holotype,  LACM 
1921.  Wreck  Bay,  Isla  San  Cristobal.  Length  16.8  mm.  FIGURE  12.  Notoacmea  rothi.  LACM  54774.  East  side,  Isla  Fernandina. 
Length  11.5  mm.  FIGURE  13.  Notoacmea  rothi.  LACM  54777.  Bahia  Cartago,  Isla  Isabela.  Length  8.8  mm.  FIGURE  14. 
Notoacmea  immaculata  new  species.  Holotype,  LACM  1923.  Isla  Baltra.  Length  5.6  mm.  FIGURE  15.  Notoacmea  immaculata. 
AHF  173-34.  Isla  Baltra.  Length  7.8  mm.  FIGURE  16.  Notoacmea  filosa  (Carpenter,  1865).  LACM  54775.  Isla  San  Cristobal. 
Length  15.5  mm. 


cave,  rounding  to  pointed  cusps.  Second  pair  of 
lateral  teeth  lateral  to  first  pair,  both  edges  con- 
vex, broad,  with  pronounced  pointed  cusps; 
third  lateral  teeth  lateral  to  second  pair,  re- 
duced, medial  edges  convex,  lateral  edges 
straight,  cusps  angular.  Marginal  teeth  lacking. 


First  lateral  plates  subrectangular;  second  lat- 
eral plates  rounded,  separated  from  third  lateral 
plates  by  partial  suture;  third  lateral  plates  bi- 
formed,  posterior  section  rounded,  lateral  sec- 
tion pointed,  extending  to  lateral  edges  of  ven- 
tral plates.  Ventral  plates  closely  set, 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


331 


subrectangular,  with  weak  anterolateral  exten- 
sions. 

Animal:  Body  pigmentation  lacking;  snout 
with  oral  lappets. 

HOLOTYPE  DIMENSIONS. — Length  16.8,  width 
14.0,  height  4.8  mm. 

TYPE-LOCALITY. — Ecuador:  Galapagos  Is- 
lands; Isla  San  Cristobal,  Wreck  Bay  (0°54'S, 
89°36'W).  Leg.  J.  DeRoy,  12  May  1968,  4  spec- 
imens. 

TYPE-MATERIAL.— Holotype,  LACM  1921 
(shell  and  radular  slide),  1  paratype,  LACM 
1922;  paratypes  also  deposited  in  the  collections 
of  CAS  and  USNM. 

DISTRIBUTION. — Galapagos  Islands;  Isla  Fer- 
nandina  (ANSP  152554),  Isla  Isabela  (SU  239), 
Isla  Rabida  (LACM  71-69),  Isla  Bartolome 
(AMNH  163290),  Isla  Santa  Cruz  (ANSP 
154889),  Isla  Baltra  (AMNH  163263),  Isla  Santa 
Maria  (CAS  23025),  Isla  Santa  Fe  (AHF  48-33), 
Isla  Espanola  (USNM  102359),  Isla  San  Cristo- 
bal (MCZ  205068). 

MATERIAL  EXAMINED. — 40  lots,  435  speci- 
mens, 9  radula  preparations. 

ETYMOLOGY. — We  are  pleased  to  name  the 
species  in  honor  of  Barry  Roth  of  the  California 
Academy  of  Sciences  in  recognition  of  his  work 
in  molluscan  systematics. 

Notoacmea  immaculata  new  species 

(Figures  14,  15,  26,  32) 

Shell  (Figs.  14-15):  Small  (maximum  length 
12  mm),  thin,  diaphanous,  height  medium.  Apex 
anterior  to  center,  anteriorly  directed;  all  slopes 
convex.  Aperture  ovoid;  sides  straight,  sides 
elevated.  Sculpture  of  faint  gray,  broad  riblets 
and  concentric  growth  lines.  Exterior  light  gray, 
mottled  with  yellow-brown,  brown,  and  white; 
darker  markings  concentrated  into  broad  pos- 
terior ray  bordered  with  white.  Interior  margin 
broad,  dull,  marked  with  exterior  pattern;  inter- 
mediate area  translucent,  glossy  white;  exterior 
pattern  readily  visible  through  shell;  central  area 
glossy,  translucent,  marked  with  sparse  yellow 
streaks,  central  stain  lacking. 

Radula  (Figs.  26,  32):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
medial  edges  convex,  lateral  edges  straight  to 
slightly  concave,  tapering  to  rounded  cusps; 
second  pair  of  lateral  teeth  posterior  and  slightly 
lateral  to  first  pair,  both  edges  convex,  tapering 
to  rounded  cusps.  First  and  second  lateral  teeth 


approximately  equal  in  width.  Third  pair  of  lat- 
eral teeth  posterior  and  lateral  to  second  pair, 
medial  edges  convex,  lateral  edges  elongate, 
straight  to  slightly  concave,  extending  to  edges 
of  ventral  plates,  cusps  rounded.  Marginal  teeth 
lacking.  First  lateral  plates  ovoid;  second  lateral 
plates  rounded  posteriorly,  separated  from  third 
lateral  plates  by  partial  suture;  third  lateral 
plates  lobate.  Ventral  plates  closely  set,  subrect- 
angular with  strong  anterior  sutures. 

Animal:  Body  pigmentation  lacking;  snout 
with  oral  lappets. 

HOLOTYPE  DIMENSIONS. — Length  5.6,  width 
4.3,  height  1.4  mm. 

TYPE-LOCALITY. — Ecuador:  Galapagos  Is- 
lands; Isla  Baltra  (0°26'S,  90°17'W),  Caleta  del 
Norte,  0-3  m.  Leg.  ANTON  BRUUN,  cr.  18B, 
sta.  791,  21  Sep.  1966,  1  specimen. 

TYPE-MATERIAL.— Holotype,  LACM  1923 
(shell  and  radula  slide),  1  paratype,  CAS  15920 
(shell  and  radula  slide).  Paratype  from  Isla  Santa 
Cruz,  Academy  Bay. 

DISTRIBUTION. — Galapagos  Islands;  Isla  Fer- 
nandina  (LACM  62-196),  Isla  Isabela  (LACM 
71-70),  Isla  Bartolome  (AMNH  163290),  Isla 
Santa  Cruz  (ANSP  154889),  Isla  Baltra  (LACM 
66-206),  Isla  San  Cristobal  (ANSP  153328). 

MATERIAL  EXAMINED. — 14  lots,  63  speci- 
mens, 5  radula  preparations. 

ETYMOLOGY. — The  name  is  a  Latin  adjective, 
immaculatus  (unstained),  referring  to  the  lack 
of  a  central  stain  in  the  area  within  the  myo- 
stracum. 

DISCUSSION. — The  radular  difference  that  is 
the  chief  basis  of  the  separation  of  the  two 
species  is  unmistakable  and  qualitative:  in  N. 
rothi  the  third  lateral  teeth  are  reduced  (Fig.  25) 
and  in  TV.  immaculata  the  third  lateral  teeth  are 
large  (Fig.  26).  The  lateral  plate  morphologies 
are  correspondingly  different.  In  N.  rothi  the 
second  and  third  lateral  plates  are  approximately 
equal  in  size,  and  the  lateral  edges  of  the  third 
lateral  plates  form  small  pointed  projections.  In 
N.  immaculata  the  third  lateral  plates  are  larger 
than  the  second  lateral  plates  and  the  lateral  pro- 
jections are  rounded. 

The  shells  of  N.  rothi  and  N.  immaculata 
have  similar  overall  proportions  and  sculpture. 
The  color  pattern  consists  of  radiating  and  scat- 
tered whitish  tessellations,  with  the  greatest 
concentration  of  white  tessellations  in  two  lat- 
ero-posterior  rays,  the  posterior  area  between 


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the  two  rays  having  the  least  amount  of  tessel- 
late  flecking  so  that  it  may  appear  to  be  a  single, 
uniformly  dark  posterior  ray.  Those  specimens 
confirmed  on  radular  examination  to  have  the 
tooth  pattern  of  N.  rothi  have  the  dark  gray- 
green  ground  color  predominating,  whereas 
those  identified  as  N.  immaculata  have  a  light 
gray  or  white  ground  color.  The  largest  speci- 
mens examined  have  proven  to  be  N.  rothi;  the 
largest  specimen  verified  as  N.  immaculata  is 
12  mm  in  length.  The  large  specimens  of  N.  rothi 
have  a  dark  interior  stain,  which  is  generally 
lacking  in  N.  immaculata.  One  small,  stunted 
specimen  verified  as  N.  immaculata  (LACM  71- 
48)  shows  a  slight  trace  of  brown  interior  stain. 
It  is  possible  that  the  brown  stain  is  indicative 
of  the  attainment  of  size  rather  than  a  species- 
specific  character.  Too  few  specimens  verified 
as  N.  immaculata  are  available  to  enable  us  to 
be  certain  that  any  shell  characters  may  be  used 
as  proof  of  identity. 

Of  the  mainland  acmaeid  species,  the  No- 
toacmea  rothi-immaculata  complex  resembles 
Notoacmea  filosa  (Carpenter,  1865)  (Fig.  6), 
which  has  similar  shell  characters.  They  differ 
from  N.  filosa  in  the  following  ways:  N.  rothi- 
immaculata  has  a  profile  of  medium  height;  N. 
filosa  has  a  low  profile.  In  N.  rothi-immaculata 
the  interspaces  are  broader  than  the  riblets;  in 
N.  filosa  the  riblets  are  more  numerous  and  the 
interspaces  approximately  equal  in  width  to  the 
riblets.  Notoacmea  filosa  has  a  color  pattern  of 
radiating  dark  and  lighter  rays,  often  interrupt- 
ed, but  not  tessellated  in  circular  or  oval  pat- 
terns. The  tessellate  markings  are  characteristic 
of  N.  rothi-immaculata.  The  dark  posterior  ray 
of  N.  rothi-immaculata  is  not  a  feature  of  N. 
filosa . 

Although  the  configuration  of  the  lateral  teeth 
of  N.  filosa  has  little  in  common  with  that  of  N. 
immaculata,  there  is  a  similarity  between  N.  fi- 
losa and  N.  rothi.  However,  the  shape  of  the 
second  lateral  teeth  differs:  in  N.  filosa  the  sec- 
ond lateral  teeth  are  triangular;  in  N.  rothi  the 
second  lateral  teeth  are  broad  with  convex 
edges. 

No  detailed  observations  on  the  habitat  of 
either  species  are  available  to  us.  We  know  from 
collection  data  on  museum  specimens  that  N. 
rothi  occurs  intertidally.  Specimens  are  relative- 
ly free  of  encrustations  except  for  some  coralline 
algae  and  spirorbid  worm  tubes.  The  edges  of 


the  apertures  are  smooth  and  oval,  not  molded 
to  fit  a  habitual  site  of  attachment,  suggesting 
that  the  normal  habitat  is  likely  to  be  on  the 
undersides  of  stones  in  tidepools.  Station  data 
for  the  holotype  of  N.  immaculata  indicate  a 
depth  of  0  to  3  m.  The  absence  of  specimens  in 
the  intertidal  collections  of  J.  DeRoy  suggests 
that  N.  immaculata  is  essentially  a  subtidal 
species. 

The  elongated  teeth  of  N.  rothi  are  similar  to 
those  of  such  temperate  species  as  Collisella 
pelta  (Rathke,  1833),  Notoacmea  persona 
(Rathke,  1833),  and  Lottia  gigantea.  All  have 
pointed  cusps  on  the  first  and  second  laterals 
and  reduced  third  laterals.  These  temperate 
species  are  known  to  feed  upon  sessile  diatoms 
and  noncalcareous  algae  in  the  middle  and  high 
intertidal  zones,  so  we  infer  that  N.  rothi  does 
also. 

The  short  blunt  teeth  of  N.  immaculata  are 
similar  to  those  of  species  known  to  feed  on  cor- 
alline algae.  The  presumed  subtidal  occurrence 
of  N.  immaculata  is  in  accordance  with  the 
abundant  subtidal  occurrence  of  coralline  algae. 

Lottia  mimica  new  species 

(Figures  17-19,  27,  33) 

Shell  (Figs.  17-19):  Size  medium  (maximum 
length  25  mm),  height  medium;  apex  anterior  to 
center;  all  slopes  convex.  Aperture  ovoid,  lat- 
eral edges  somewhat  parallel.  Sculpture  of 
raised  angular  ribs  with  one  or  two  secondary 
ribs  between  each  pair  of  primary  ribs;  ribs  ex- 
tending slightly,  crenulating  the  margin.  Exte- 
rior gray-brown  with  white  radial  markings  that 
may  or  may  not  correspond  to  ribs.  Apex  white, 
with  fine,  dark  radial  lines  typically  concentrat- 
ed in  rays.  Interior  margin  dark,  with  white 
markings  corresponding  to  exterior  color  pat- 
tern; intermediate  area  blue- white;  central  area 
stained  with  dark  brown;  apical  region  white; 
exterior  markings  visible  through  shell. 

Radula  (Figs.  27,  33):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
medial  edges  convex,  lateral  edges  straight, 
cusps  pointed.  Second  pair  of  lateral  teeth  po- 
sitioned posterior  to  and  slightly  lateral  to  first 
pair,  medial  edges  convex,  lateral  edges  slightly 
convex,  cusps  pointed.  Third  lateral  teeth  lateral 
to  second  pair,  medial  edges  strongly  convex, 
lateral  edges  concave,  cusps  pointed.  All  lateral 
teeth  approximately  equal  in  width.  Marginal 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


333 


FIGURES  17-22.  New  species  of  Lottia  from  the  Galapagos  Islands.  All  from  Academy  Bay,  Isla  Santa  Cruz.  FIGURE  17. 
Lottia  mimica  new  species.  Holotype,  LACM  1925.  Length  16.2  mm.  FIGURE  18.  Lottia  mimica.  Paratype,  LACM  1926. 
Length  14.8  mm.  FIGURE  19.  Lottia  mimica.  Paratype,  LACM  1926.  Length  9.6  mm.  FIGURE  20.  Lottia  smithi  new  species. 
Holotype,  LACM  1927.  Length  12.4  mm.  FIGURE  21.  Lottia  smithi.  Paratype,  LACM  1928.  Length  12.9  mm.  FIGURE  22. 
Lottia  smithi.  Paratype,  LACM  1928.  Length  7.5  mm. 


teeth  small,  narrow,  extending  over  ventral 
plates  in  vicinity  of  third  pair  of  lateral  teeth. 
First  lateral  plates  square,  slightly  overlapping 
anterior  ribbon  segments;  second  lateral  plates 
rounded,  separated  from  third  lateral  plates  by 
partial  suture;  third  lateral  plates  irregular,  with 


prominent  lateral  extensions  extending  to  edges 
of  ventral  plates.  Ventral  plates  closely  set,  with 
broad,  rounded  anterior  process,  lateral  edges 
concave,  posterior  process  weak. 

Animal:     Base  of  every  second  or  third  mantle 
tentacle  with  dark  red-brown  pigmentation; 


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snout  with  oral  lappets;  secondary  gill  complete 
but  reduced  in  front  of  head,  composed  of  lap- 
pets (approximately  14  per  mm);  every  other 
lappet  reduced,  less  than  one-half  the  size  of  the 
larger  ones. 

HOLOTYPE  DIMENSIONS. — Length  16.2,  width 
12.4,  height  4.2  mm. 

TYPE-LOCALITY. — Ecuador:  Galapagos  Is- 
lands; Isla  Santa  Cruz  (0°38'S,  90°23'W),  Acad- 
emy Bay,  Coamano  Island,  intertidal  zone.  Leg. 
J.  DeRoy,  Oct.  1967. 

TYPE-MATERIAL.— Holotype,  LACM  1925 
(shell  and  radula  slide),  17  paratypes,  LACM 
1926,  paratypes  also  deposited  in  the  collections 
of  CAS  and  USNM.  Paratypes  collected  at  sev- 
eral stations  in  Academy  Bay  by  J.  DeRoy  be- 
tween 1967  and  1969. 

DISTRIBUTION. — Galapagos  Islands;  Isla  Fer- 
nandina  (AMNH  163363),  Isla  Isabela  (AHF  74- 
33),  Isla  Bartolome  (AMNH  163290),  Isla  Santa 
Fe  (AMNH  163362),  Isla  Espanola  (AHF  359- 
35),  and  Isla  San  Cristobal  (CAS  23103). 

MATERIAL  EXAMINED. — 33  lots,  198  speci- 
mens, 23  radula  preparations. 

ETYMOLOGY. — The  name  mimica  is  a  Latin 
adjective,  imitative,  indicative  of  the  difficulty 
of  distinguishing  the  two  members  of  the  Lottia 
pair  by  their  external  appearance. 

Lottia  smith!  new  species 

(Figures  20-22,  28,  34) 

Shell  (Figs.  20-22):  Size  medium  (maximum 
length  25  mm),  height  medium;  apex  positioned 
in  anterior  third  of  shell;  all  slopes  convex.  Ap- 
erture ovoid.  Sculpture  of  rounded  primary  ribs, 
secondary  ribs  of  equal  strength  but  beginning 
below  apex.  Primary  and  secondary  ribs  white, 
interspaces  brown;  apex  white,  with  fine  dark 
radial  lines  gathered  into  rays;  interior  margin 
dull  yellow  with  irregular  brown  markings  that 
correspond  to  exterior  interspaces;  intermediate 
area  and  central  areas  white;  interior  of  myo- 
stracum  bordered  by  yellow-brown  halo.  Exte- 
rior markings  visible  through  shell. 

Radula  (Figs.  28,  34):  First  pair  of  lateral  teeth 
closely  set  at  anterior  edge  of  ribbon  segment, 
both  edges  convex,  rounding  to  blunt  cusps; 
second  pair  of  lateral  teeth  posterior  and  slightly 
lateral  to  first  pair,  both  edges  convex,  rounding 
to  blunt  cusps.  Third  pair  of  lateral  teeth  posi- 
tioned lateral  and  posterior  to  second  pair,  both 
edges  convex,  rounding  to  blunt  cusps.  Marginal 


teeth  small,  narrow,  overlapping  ventral  plates 
just  anterior  of  third  pair  of  lateral  teeth.  First 
lateral  plates  ovoid,  slightly  overlapping  anterior 
ribbon  segment;  second  lateral  plates  distinctly 
smaller  than  other  lateral  plates,  medial  edges 
rounded,  separated  from  third  lateral  plates  by 
a  partial  suture.  Posterior  edge  of  third  lateral 
plates  concave,  with  lateral  extensions  termi- 
nating in  strongly  hooked  edges.  Ventral  plates 
closely  set  with  strong  posterior  process;  ante- 
rior process  also  present.  Lateral  edges  in  vicin- 
ity of  marginal  teeth  concave;  anterior  sutures 
parallel  with  anterior  edges  of  ventral  plates. 

Animal:  Mantle  tentacle  pigmentation  some- 
times present;  snout  with  oral  lappets;  second- 
ary gill  complete,  but  reduced  in  front  of  head, 
composed  of  lappets  (approximately  11  per 
mm). 

HOLOTYPE  DIMENSIONS. — Length  12.4  mm, 
width  9.7  mm,  height  4.5  mm. 

TYPE-LOCALITY. — Ecuador:  Galapagos  Is- 
lands; Isla  Santa  Cruz  (0°38'S,  90°23'W),  Acad- 
emy Bay,  Punta  Nunez,  intertidal  zone.  Leg.  J. 
DeRoy,  13  Oct.  1969. 

TYPE-MATERIAL.— Holotype,  LACM  1927 
(shell  and  radula  slide),  9  paratypes,  LACM 
1928;  paratypes  also  deposited  in  the  collections 
of  CAS  and  USNM.  All  type-material  from  Isla 
Santa  Cruz,  Academy  Bay. 

DISTRIBUTION. — Galapagos  Islands;  Isla  Fer- 
nandina  (LACM  72-196),  Isla  Isabela  (CAS 
27221),  Isla  Bartolome  (AMNH  163290),  Isla 
Santa  Cruz  (LACM  28839),  Isla  Santa  Maria 
(ANSP  153370),  Isla  San  Cristobal  (ANSP 
153328). 

MATERIAL  EXAMINED. — 15  lots,  70  speci- 
mens, 14  radula  preparations. 

ETYMOLOGY. — We  are  pleased  to  name  this 
species  in  honor  of  the  late  Allyn  G.  Smith  of 
the  California  Academy  of  Sciences  in  recogni- 
tion of  his  work  with  eastern  Pacific  mollusks, 
including  those  of  the  Galapagos  Islands. 

DISCUSSION. — The  radular  difference  that 
separates  L.  mimica  and  L.  smithi  is  readily  ap- 
parent. In  L.  mimica  the  lateral  teeth  are  point- 
ed distally;  in  L.  smithi  they  are  rounded.  The 
third  lateral  teeth  of  L.  mimica  are  of  the  same 
width  as  the  second;  in  L.  smithi  the  third  lateral 
teeth  are  much  broader  than  the  second.  Lottia 
mimica  lacks  the  strong  posterior  process  on  the 
ventral  plates  that  is  present  in  L.  smithi. 

In  addition  to  radular  difference,  there  is 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


335 


• 


.  25 


27 


FIGURES  23-28.  Radular  dentition.  FIGURE  23.  Notoacmea  uhiquita  new  species.  FIGURE  24.  Notoacmea  pumila  new 
species.  FIGURE  25.  Notoacmea  rothi  new  species.  FIGURE  26.  Notoacmea  immaculata  new  species.  FIGURE  27.  Lottia  mimica 
new  species.  FIGURE  28.  Lottia  smithi  new  species.  Anterior  towards  top  of  page. 


another  significant  anatomical  difference:  in  L. 
mimica  the  mantle  tentacles  are  darkly  pig- 
mented  (Fig.  3)  in  much  the  same  way  as  in  L. 
gigantea  (Fig.  1).  This  pigmentation  is  weakly 
developed  or  entirely  lacking  in  L.  smithi.  This 
distinction  could  prove  useful  in  future  field 
studies  because  it  provides  a  reliable,  nonfatal 
method  of  species  determination. 

Shells  of  L.  mimica  and  L.  smithi  are  essen- 
tially indistinguishable  and  are  characterized  by 
moderately  strong  radial  ribs,  variable  in  the 
number  reaching  the  margin.  Some  specimens 
of  both  species  have  relatively  few  primary  and 
secondary  ribs,  and  these  ribs  project,  crenulat- 
ing  the  margin  (Figs.  18,  21).  In  others  the  sec- 
ondary ribs  are  more  numerous  and  the  primary 
ribs  less  prominent.  In  these  specimens  the  ribs 
project  only  slightly  and  the  shell  margin  is  rel- 
atively even  (Figs.  17,  19,  20,  22).  The  normal 
pattern  on  the  apical  region  of  juvenile  shells  is 
identical  in  both  species  (Figs.  19-22).  The  api- 
cal tip  is  dark  colored  with  a  pattern  of  thin, 
dark  lines,  concentrated  in  six  bundles  in  the  1, 


3,  5,  7,  9,  and  1 1  o'clock  positions.  In  most  spec- 
imens this  pattern  changes  abruptly  to  one  in 
which  rib  surfaces  are  lighter  colored  and  the 
interspaces  darker,  often  showing  concentric 
variations  in  intensity.  In  a  few  specimens  of 
both  species  (Figs.  19,  22),  the  juvenile  pattern 
changes  to  a  solid  6-rayed  pattern  in  the  adult. 
The  holotype  of  L.  mimica  is  unusual;  in  the 
early  stage  it  is  uniformly  dark,  changing  abrupt- 
ly to  a  rayed  pattern  in  which  the  lighter  rays  do 
not  necessarily  correspond  to  the  ribs.  Four 
specimens  of  the  type  lot  of  L.  mimica,  includ- 
ing the  holotype,  are  predominantly  dark  col- 
ored; none  in  the  type  lot  of  L.  smithi  may  be 
so  described.  The  range  of  variability  of  L.  mim- 
ica is  therefore  somewhat  broader  than  that  of 
L.  smithi. 

The  peculiar  markings  in  the  juvenile  shell  of 
L.  mimica-smithi  as  well  as  its  particular  adult 
pattern  are  unlike  those  of  any  other  acmaeid 
species.  The  extreme  specimens  with  few  ribs 
are  similar  to  Collisella  pediculus,  although  that 
species  has  fewer,  more  prominent  ribs.  Co///- 


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29 


30 


31 


32 


33 


34 


FIGURES  29-34.  Radular  basal  plates.  FIGURE  29.  Notoacmea  ubiquita  new  species.  FIGURE  30.  Notoacmea  pumila  new 
species.  FIGURE  31.  Notoacmea  rothi  new  species.  FIGURE  32.  Notoacmea  immaculata  new  species.  FIGURE  33.  Lottia  mimica 
new  species.  FIGURE  34.  Lottia  smith!  new  species.  Anterior  towards  top  of  page. 


sella  mitella  has  a  greater  number  of  ribs  and 
has  a  more  uniform  color  pattern,  with  lighter 
ribs  and  darker  interspaces. 

There  is  no  particular  similarity  to  the  large, 
dark  brown  shells  of  Lottia  gigantea  or  the  blue- 
green  shells  of  L.  mesoleuca  and  L.  stipulata, 
but  that  is  not  surprising  in  view  of  the  lack  of 
consistent  shell  characters  within  all  acmaeid 
genera. 


We  have  no  information  about  the  habitat  of 
either  L.  mimica  or  L.  smithi.  The  original  col- 
lecting information  supplied  by  J.  DeRoy  indi- 
cates that  they  were  collected  on  rocks  exposed 
to  heavy  surf.  All  of  the  shells  in  both  type  lots 
were  heavily  encrusted  with  coralline  algae,  as 
are  those  of  the  Panamic  species  of  Collisella 
that  live  under  exposed  surf  conditions,  such  as 
C.  pediculus.  Limpets  in  this  habitat  have  an 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE 


337 


habitual  site  of  attachment  with  the  shell  edge 
molded  to  fit  the  home  site.  The  margins  of  L. 
mimica  and  L.  smithi  are  sufficiently  irregular 
to  suggest  that  they  conform  to  specific  sites. 

The  short,  blunt,  equal-sized  radular  teeth  of 
L.  smithi  are  well  adapted  for  feeding  on  cor- 
alline algae.  The  shape  of  the  lateral  teeth,  par- 
ticularly the  expansion  of  the  third  laterals,  is 
similar  to  that  found  in  other  coralline-feeding 
Collisella  of  the  tropical  eastern  Pacific.  The  lat- 
eral teeth  of  L.  mimica,  however,  are  not  those 
of  a  coralline  feeder,  nor  are  they  like  those  of 
most  of  the  diatom-feeding  species  of  Notoac- 
mea  or  Collisella,  in  which  the  third  lateral  teeth 
are  reduced.  The  medium  length  teeth  with 
pointed  cusps,  all  equally  large,  are  like  those 
of  many  of  the  Peruvian  Scurria  and  the  Cali- 
fornian  Notoacmea  insessa  (Hinds,  1842).  The 
last  species  is  known  to  feed  on  the  stipes  of 
brown  algae.  The  teeth  of  L.  mimica  are  most 
likely  adapted  for  feeding  on  some  of  the  fleshy, 
encrusting,  but  noncalcareous  algae. 

Limpets  of  the  L.  mimica— smithi  type  are 
present  in  late  Pleistocene  deposits  on  Isla  San 
Salvador  (Hertlein  and  Strong  1939).  The  range 
of  variation  in  the  fossil  specimens  is  similar  to 
that  seen  in  Recent  specimens  of  both  L.  mimica 
and  L.  smithi.  However,  specific  identifications 
are  not  possible  from  the  shells  alone. 

ACMAEIDAE  OF  THE  GALAPAGOS  ISLANDS 

The  Galapagos  Islands,  or  Archipielago  de 
Colon,  are  located  approximately  800  km  west 
of  Cabo  San  Lorenzo,  Ecuador.  The  fifteen  is- 
lands and  numerous  islets  extend  from  1°40'N 
to  1°36'S  and  from  89°17'W  to  90°01'W.  The  is- 
lands have  been  the  subject  of  numerous  sci- 
entific explorations  (see  Slevin  1959)  and  are  re- 
nowned for  their  unique  fauna  and  flora.  The 
marine  molluscan  fauna  comprises  tropical  east- 
ern Pacific  elements,  endemics,  and  a  few  forms 
from  the  Indo- Pacific  faunal  region  (Emerson 
1978). 

Some  nine  different  taxa  of  acmaeid  limpets 
have  been  reported  from  the  Galapagos  Islands 
since  1855  (Reeve  1855;  Carpenter  1864;  Wim- 
mer  1880;  Stearns  1893;  Pilsbry  and  Vanatta 
1902;  Dall  1909;  Schwengel  1938;  Hertlein  and 
Strong  1955;  Keen  1958;  McLean  1971).  None 
of  the  species  reported  have  been  recognized  as 
endemics,  having  been  considered  instead  as 
vagrants  from  the  Calif ornian,  Panamic,  and  Pe- 


ruvian molluscan  faunal  provinces.  Many  of  the 
records  have  been  based  on  beachworn  shells  or 
small  shells  dredged  dead  and  in  poor  condition. 
Such  specimens  are  difficult  to  refer  to  known 
species  and  could  hardly  be  recognized  as  dis- 
tinct new  species.  After  examining  a  number  of 
museum  collections,  we  are  able  to  confirm  the 
presence  of  only  two  previously  reported 
species  at  the  Galapagos  Islands,  Notoacmea 
filosa  and  Lottia  mesoleuca. 

Specimens  of  L.  mesoleuca  from  the  Gala- 
pagos were  originally  misidentified  as  the  Aus- 
tralian species  Patelloida  striata  Quoy  and  Gai- 
mard,  1834,  and  were  so  treated  by  Reeve 
(1855:pl.  33,  fig.  99)  (as  "Patella  striata"),  Car- 
penter (1864)  (as  "Acmaea  striata  Reeve"),  and 
Stearns  (1893)  (as  "Acmaea  striata  Reeve"). 
Occurrences  of  L.  mesoleuca  at  the  Galapagos 
are  rare.  It  is  represented  in  the  collections  ex- 
amined by  three  specimens:  two  from  Isla 
Genovesa  (AHF  782-38)  and  a  single  beach 
specimen  from  the  "Galapagos"  (ANSP  39189). 
Isla  Genovesa  is  the  most  northeastern  of  the 
islands,  and  it  is  conceivable  that  the  species 
may  be  established  there  and  not  elsewhere  in 
the  archipelago. 

Notoacmea  filosa  is  also  rare  in  collections. 
It  is  represented  by  only  two  museum  lots,  one 
from  Isla  Santa  Cruz  (AMNH  177320)  and  one 
from  Isla  San  Cristobal  (LACM  54775)  (Fig.  16) 
collected  in  1929.  Further  information  about  the 
occurrence  of  these  two  species  at  the  Galapa- 
gos Islands  is  desirable.  Considering  that  recent 
collecting  efforts  have  not  produced  these 
species,  we  only  provisionally  list  them  in  the 
Galapagos  Islands  fauna. 

We  consider  all  other  records  of  Acmaeidae 
from  the  Galapagos  Islands  to  be  misidentifica- 
tions  of  the  four  new  species  described  herein. 
Because  separation  of  members  of  the  species 
pairs  is  based  on  radular  characters,  it  is  not 
possible  to  list  species-specific  synonymies  for 
each  member. 

In  both  species  pairs,  additional  characters 
segregate  with  the  radular  morphotypes.  In  the 
Notoacmea  siblings,  there  are  differences  in 
shell  size  and  coloration  associated  with  the  rad- 
ular types.  In  the  Lottia  pair,  there  are  no  sig- 
nificant differences  in  shell  size,  sculpture,  or 
coloration;  however,  there  are  differences  in 
mantle  pigmentation  and  secondary  gill  mor- 
phology. We  interpret  these  separate  character 


338 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  12 


states  as  evidence  against  a  simple  radular  poly- 
morphism and  instead  recognize  the  pairs  as 
separate  species.  There  is  also  evidence  that  the 
habitat  differs  in  the  Notoacmea  pair  but  not  in 
the  Lottia  pair. 

Whether  speciation  of  the  two  pairs  of  sibling 
species  was  sympatric  or  allopatric  is  unknown. 
However,  we  hope  that  future  workers  will  pur- 
sue this  problem  in  the  field  and  the  laboratory. 
Electrophoretic  analysis  could  prove  produc- 
tive. 

ACKNOWLEDGMENTS 

We  are  indebted  to  Jacqueline  DeRoy  of  Isla 
Santa  Cruz,  Galapagos  Islands,  who  furnished 
preserved  specimens  of  the  new  species  from 
those  islands.  Donald  R.  Shasky  of  Redlands, 
California,  loaned  pertinent  specimens  from  his 
collection.  Courtesies  were  extended  to  Lind- 
berg  on  his  museum  visits  by  George  M.  Davis 
(ANSP),  William  K.  Emerson  (AMNH),  Welton 
L.  Lee  (CAS),  Joseph  Rosewater  and  Kathy 
Lamb  (USNM),  Barry  Roth  (CAS),  and  Ruth 
Turner  (MCZ).  Barry  Roth  also  made  available 
the  late  Leo  G.  Hertlein's  unpublished  notes  on 
the  Galapagos  molluscan  fauna.  Bertram  C. 
Draper,  Los  Angeles,  made  the  prints  of  shell 
specimens,  and  Sally  Walker,  University  of  Cal- 
ifornia, Santa  Cruz,  prepared  the  line  drawing. 
We  thank  Eugene  V.  Coan,  Myra  Keen,  and 
Barry  Roth  for  reading  the  manuscript  and  of- 
fering helpful  suggestions. 

RESUMEN 

La  definition  de  genero  esta  basada  en  las  ca- 
racteristicas  conservativas  de  la  estructura  de  la 
concha  y  tambien  en  las  caracteristicas  cualita- 
tivas  de  la  radula.  La  Lottia,  que  anteriormente 
se  habia  considerado  monotfpica,  se  extende  a 
incluir  las  especies  panamicas  con  la  agalla  se- 
cundaria  (un  cordon  branquial),  que  anterior- 
mente se  habia  atribuido  a  la  Scurria.  La  Scur- 
ria  tiene  la  agalla  semejante,  pero  la  estructura 
de  la  concha  se  diferencia.  Las  nuevas  especies 
Notoacmea  ubiquita  de  Mexico  y  N.  pumila  de 
Ecuador  tienen  las  conchas  pequenas  y  son  es- 
pecies alopatricas  con  los  dientes  radulares  que 
estan  modificados  para  alimentarse  de  la  alga 
coralina.  Dos  nuevas  especies  de  Notoacmea 
(N.  rothi  y  N.  immaculata),  que  son  endemicas 
a  las  Islas  Galapagos,  constituyen  un  par  de  es- 
pecies que  se  diferencian  principalmente  por  los 


rasgos  de  la  radula:  los  dientes  radulares  de  N. 
immaculata  estan  adoptados  para  alimentarse 
de  la  alga  calcarea;  los  dientes  radulares  de  N. 
rothi,  para  alimentarse  de  la  alga  no  calcarea. 
Un  par  de  nuevas  especies  de  Lottia  endemicas 
a  las  Islas  Galapagos  (L.  mimica  y  L.  smithi) 
tambien  se  diferencian  principalmente  por  las 
caracteristicas  radulares.  La  Lottia  mimica  se 
alimenta  de  la  alga  y  es  no  calcarea;  y  la  L.  smithi 
se  alimenta  de  alga  y  es  calcarea.  Estas  cuatro  es- 
pecies endemicas  son  las  principales  lapas  ac- 
maeidas  de  las  Islas  Galapagos.  Dos  especies  de 
la  tierra  firme,  Notoacmea  filosa  y  Lottia  me- 
soleuca,  se  han  observado  no  mas  esporadica- 
mente  en  las  Islas  Galapagos. 

LITERATURE  CITED 

CARPENTER,  P.  P.  1864.  Supplementary  report  on  the  present 
state  of  our  knowledge  with  regard  to  the  Mollusca  of  the 
west  coast  of  North  America.  Rep.  Brit.  Assoc.  Adv.  Sci. 
1864:517-686. 

CHRISTIAENS,  J.  1975.  Revision  provisoire  des  mollusques 
marins  recents  de  la  famille  des  Acmaeidae.  Inf.  Soc.  Beige 
Malacol.  4(4):3-20. 

DALL,  W.  H.  1871.  On  the  limpets;  with  special  reference  to 
the  species  of  the  west  coast  of  America,  and  to  a  more 
natural  classification  of  the  group.  Am.  J.  Conchol. 
6(3):227-282. 

.  1909.  Report  on  a  collection  of  shells  from  Peru,  with 

a  summary  of  the  littoral  marine  Mollusca  of  the  Peruvian 
zoological  province.  Proc.  U.S.  Natl.  Mus.  37:147-294. 

EMERSON,  W.  K.  1978.  Mollusks  with  Indo-Pacific  faunal 
affinities  in  the  eastern  Pacific  Ocean.  Nautilus  92:91-96. 

HERTLEIN,  L.  G.,  AND  A.  M.  STRONG.  1939.  Marine  Pleis- 
tocene mollusks  from  the  Galapagos  Islands.  Proc.  Calif. 
Acad.  Sci.,  ser.  4,  23:367-380. 

,  AND .  1955.  Marine  mollusks  collected  at  the 

Galapagos  Islands  during  the  voyages  of  the  Velero  III, 
1931-1932.  Pp.  111-115  in  Essays  in  the  Natural  Sciences 
in  Honor  of  Capt.  Allan  Hancock,  Univ.  So.  Calif.,  Los 
Angeles,  Calif. 

KEEN,  A.  M.  1958.  Sea  shells  of  tropical  west  America.  1st 
ed.  Stanford  Univ.,  Stanford,  Calif.  624  pp. 

LINDBERG,  D.  R.  1976.  Cenozoic  phylogeny  and  zoogeog- 
raphy of  the  Acmaeidae  in  the  eastern  Pacific.  Ann.  Rep. 
West.  Soc.  Malacol.  9:15-16. 

.  1978.  On  the  taxonomic  affinities  of  Collisella  ed- 

mitchelli,  a  late  Pleistocene  limpet  from  San  Nicolas  Island, 
California.  Bull.  So.  Calif.  Acad.  Sci.  77:65-70. 

.   1979.  Variation  in  the  limpet  Collisella  ochracea  and 


the  northeastern  Pacific  distribution  of  Notoacmea  testu- 
dinalis  (Acmaeidae).  Nautilus  93:50-56. 

MACCLINTOCK,  C.  1967.  Shell  structure  of  patelloid  and  bel- 
lerophontoid  gastropods  (Mollusca).  Peabody  Mus.  Nat. 
Hist.  Yale  Univ.  Bull.  22:1-140. 

MCLEAN,  J.  H.  1966.  West  American  prosobranch  Gastro- 
poda: Superfamilies  Patellacea,  Pleurtomariacea,  and  Fissu- 
rellacea.  Ph.D.  dissertation,  Stanford  Univ.,  Stanford,  Cal- 
if. 255  pp. 


LINDBERG  &  McLEAN:  TROPICAL  EASTERN  PACIFIC  ACMAEIDAE  339 

.   1971.  Family  Acmaeidae,  in  Keen,  A.  M.,  Sea  shells  SCHWENGEL,  J.  S.    1938.    Zoological  results  of  the  George 

of  tropical  west  America.  2nd  ed.  Stanford  Univ.  Press,  Vanderbilt  South  Pacific  Expedition,  1937.  Part  I.  Galapa- 

Stanford,  Calif.  1064  pp.  gos  Mollusca.  Proc.  Acad.  Nat.  Sci.  Philadelphia  90:1-3. 

1973.  Family  Acmaeidae,  in  Marincovich,  L.,  Inter-  SLEVIN.J.  R.   1959.  The  Galapagos  Islands:  A  history  of  their 


tidal  marine  mollusks  of  Iquique,  Chile.  Nat.  Hist.  Mus.  exploration.  Occas.  Pap.  Calif.  Acad.  Sci.  25:1-150. 

Los  Angeles  Co.  Sci.  Bull.  16:1-49.  STEARNS,  R.  E.  C.  1893.  Report  on  the  mollusk  fauna  of  the 

PILSBRY,  H.  A.,  AND  E.  G.  VANATTA.  1902.  Papers  from  the  Galapagos  Islands  with  descriptions  of  new  species.  Proc. 

Hopkins  Stanford  Galapagos  Expedition,  1898-99,  no.  13,  U.S.  Natl.  Mus.  16:353-450. 

marine  Mollusca.  Proc.  Washington  Acad.  Sci.  4:549-560.  WIMMER,  A.  1880.  Zur  Conchylien-Fauna  der  Galapagos-In- 

REEVE,  L.  1855.  Conchologia  iconica:  or,  illustrations  of  the  seln.  Sitzungsber.  K.  Akad.  Wiss.  Wien  Math.-Natwiss.  Kl. 

shells  of  molluscous  animals.  Vol.  8.  Monograph  of  the  ge-  80(Abt.  I)(10)(1879):465-514. 

nus  Patella.  London.  Pages  not  numbered,  41  pis. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  941 18 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  13,  pp.  341-348,  5  figs. 


June  24,  1981 


A  REVIEW  OF  THE  BATHYAL  FISH  GENUS  ANTIMORA 
(MORIDAE:  GADIFORMES) 

By 
Gregg  J.  Small* 

Systematics  Laboratory,  National  Marine  Fisheries  Service,  NOAA, 
National  Museum  of  Natural  History,  Washington,  DC  20560 


ABSTRACT:  There  are  two  valid  species  of  the  genus  Antimora:  A.  microlepis  Bean  in  the  North  Pacific,  and 
A.  rostrata  (Giinther)  from  the  southeastern  Pacific,  Southern  Ocean,  and  Atlantic  Ocean.  Junior  synonyms  of 
A.  rostrata  include:  A.  australis  Barnard,  A.  meadi  Pequeno,  A.  rhina  Garman,  and  Haloporphyrus  viola  Goode 
and  Bean.  Antimora  microlepis  has  90  to  103  gill  filaments  on  the  first  arch;  A.  rostrata  has  76  to  90.  Differences 
in  the  regression  equations  of  gill  filament  length  on  standard  length,  and  of  head  length  on  standard  length 
between  fish  from  several  geographic  areas  are  shown.  Other  characters  examined  include  numbers  of  vertebrae, 
fin  rays,  gill  rakers,  and  scale  rows;  and  morphometric  ratios,  using  lengths  of  eye,  snout,  predorsal,  first 
dorsal  fin  ray,  maxillary,  and  gill  rakers,  and  width  of  interorbital. 


INTRODUCTION 

Fishes  of  the  benthopelagic  morid  genus  An- 
timora are  widely  distributed  in  the  world 
oceans,  ranging  in  depth  between  402  and  2905 
m  (Grey  1956),  and  are  very  abundant  in  some 
regions  (Wenner  and  Musick  1977).  They  appear 
to  be  most  common  on  the  continental  slopes  of 
subarctic,  subantarctic,  and  temperate  regions, 
but  are  generally  rare  in  the  subtropics  and  trop- 
ics, although  they  are  apparently  common  in  the 
vicinity  of  the  Hawaiian  and  Galapagos  islands. 

The  genus  Antimora  is  distinguished  from 
other  members  of  the  family  Moridae  by  the 
combination  of  a  pronounced  pointed  snout,  a 
pelvic  fin  with  six  rays,  a  well-developed  mental 
barbel,  a  long-based  second  dorsal  fin  with  more 
than  50  rays,  and  a  deep  indentation  in  the  out- 


*  Present  address:  Northwest  and  Alaska  Fisheries  Center, 
National  Marine  Fisheries  Service,  NOAA,  2725  Montlake 
Boulevard  East,  Seattle,  Washington  98112. 


line  of  the  anal  fin  (Svetovidov  1948).  There  is 
little  information  on  food  habits  because  speci- 
mens brought  to  the  surface  routinely  evert  their 
stomachs.  Sedberry  and  Musick  (1978)  found 
only  10  specimens  with  intact  stomachs  in  nu- 
merous deep-water  trawlings.  Individuals  with 
ripe  eggs  are  unknown,  and  specimens  smaller 
than  100  mm  are  rare.  Males  longer  than  325  mm 
are  uncommon,  although  females  are  often  long- 
er than  600  mm.  Within  its  depth  range,  there  is 
a  segregation  by  sex  and  also  by  size  of  individ- 
uals (Iwamoto  1975;  Wenner  and  Musick  1977). 
Available  information  on  the  taxonomy  and  dis- 
tribution of  the  genus  was  summarized  by  Iwa- 
moto (1975). 

Although  this  study  shows  that  only  Antimora 
rostrata,  occurring  in  all  areas  except  the  North 
Pacific,  and  A.  microlepis,  occurring  only  in  the 
North  Pacific,  are  valid  species  of  the  genus 
Antimora,  the  following  species  and  their  as- 
sociated localities  have  been  previously  pro- 
posed: Haloporphyrus  rostrata  Giinther,  1878, 


[341] 


342 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  13 


FIGURE  1.     World  localities  from  which  specimens  of  Antimora  spp.  were  studied.  Stars  =  A.  microlepis,  dots  =  A.  rostrata. 
Heavy  black  lines  divide  regions. 


from  subantarctic  seas;  Haloporphyrus  viola 
Goode  and  Bean,  1878,  from  the  temperate 
western  North  Atlantic;  Antimora  microlepis 
Bean,  1890,  from  the  temperate  eastern  North 
Pacific;  Antimora  rhina  Garman,  1899,  from  the 
Gulf  of  Panama;  Antimora  australis  Barnard, 
1925,  from  off  the  Cape  of  Good  Hope;  and  An- 
timora meadi  Pequeno,  1970,  from  off  the  coast 
of  Chile. 

No  comparison  has  ever  been  made  of  popu- 
lations represented  by  these  six  names.  The 
closest  approach  was  that  of  Schroeder  (1940) 
who  examined  limited  material  of  the  first  four 
nominal  species  (he  apparently  overlooked  A. 
australis)  and  suggested  that  they  be  referred  to 
the  same  species.  The  object  of  the  present  pa- 
per is  to  resolve  the  number  of  valid  species  of 
Antimora  using  measurements,  counts,  and  the 
known  species  distributions. 

Biological  and  distributional  information  are 
useful  in  clarifying  the  taxonomy  of  Antimora. 
It  has  been  shown  that  in  the  western  North 
Atlantic  Antimora  becomes  larger  with  increas- 
ing depth  but  does  not  reproduce  within  the  area 


of  its  greatest  known  abundance  (Wenner  and 
Musick  1977).  Migrations  may  be  a  regular  part 
of  the  life  history  of  the  western  North  Atlantic 
population.  The  mobility  of  Antimora  has  been 
confirmed  by  Cohen  (1977)  who  has  shown  that 
these  fish  are  able  swimmers.  Perhaps  there  is 
a  single  interbreeding  North  Atlantic  population. 
If  Antimora  from  other  regions  have  the  same 
swimming  ability,  then  there  may  be  single  pop- 
ulations in  the  southeastern  Pacific,  the  southern 
oceans,  and  the  northern  Pacific.  Although  data 
for  this  study  were  originally  segregated  on  the 
basis  of  six  geographical  regions  (see  Materials), 
characters  of  Antimora  in  the  western  and  east- 
ern North  Pacific  were  found  to  be  similar  as 
were  those  from  the  western  and  eastern  North 
Atlantic.  A  preliminary  analysis  indicated  that 
the  four  geographical  groupings  mentioned 
above  provide  a  more  appropriate  basis  for  com- 
parison (Fig.  1). 

METHODS 

Measurements  and  counts  were  made  accord- 
ing to  Hubbs  and  Lagler  (1970)  and  include: 


SMALL:  FISH  GENUS  ANTIMORA 


343 


standard  length,  head  length,  snout  length,  pre- 
dorsal  length,  eye  diameter,  upper  jaw  length, 
interorbital  distance,  first  dorsal  fin  ray  length, 
number  of  scales  along  the  lateral  line,  and  num- 
ber of  scale  rows  between  lateral  line  and  dorsal 
origin.  Numbers  of  vertebrae  (including  the  hy- 
pural  plate),  anal  rays,  and  dorsal  rays  were  read 
from  radiographs. 

Numbers  of  gill  filaments  and  gill  rakers  on 
the  first  gill  arch,  lengths  of  the  longest  gill  raker 
and  gill  filament  (measured  from  base  to  tip,  Fig. 
2),  length  of  longest  gill  raker  at  the  gill  angle, 
and  dorsal  fin  ray  length  (from  the  anterior  base 
to  the  tip  of  the  first  ray)  were  compared.  Bro- 
ken and  otherwise  damaged  rays  were  not  mea- 
sured. 

Coloration,  often  used  in  early  descriptions, 
was  not  recorded  due  to  color  changes  which 
occur  in  preserved  specimens.  However,  an  at- 
tempt was  made  to  recheck  other  characters 
presented  in  original  descriptions. 

MATERIALS 

A  total  of  449  specimens  were  examined  (refer  to  figures  for 
length  summary). 

WESTERN  NORTH  PACIFIC  (between  33°N  to  48°N  and  135°E 
to  145°E;  6  specimens):  U.S.  National  Museum  (USNM): 
161494  (1);  160607  (1);  160606  (1);  149228  (1);  117886  (2). 

EASTERN  NORTH  PACIFIC  (between  18°N  and  56°N;  %  spec- 
imens): California  Academy  of  Sciences  (CAS):  3883  (2); 
37559  (1);  34354  (2);  34353  (2);  32308  (2);  27525  (1);  26226  (1); 
uncat.  (3);  CAS-SU  5276  (5);  77  (1).  Museum  of  Comparative 
Zoology  (MCZ):  28250  (1).  Scripps  Institution  of  Oceanogra- 
phy (SIO):  70-249  (22);  70-247  (2);  68-443  (1);  59-265  (1).  Uni- 
versity of  Washington  (UW):  19309  (2);  19235  (1);  19228  (7); 
19139  (4);  18492  (8);  18201  (2)  18190  (2);  17180  (7);  19149  (1); 
18493  (5).  USNM:  45361  (2  syntypes  of  Antimora  microlepis); 
54573  (1);  54364  (1);  53876  (3);  48562  (1);  47238  (1);  47237  (1). 

WESTERN  NORTH  ATLANTIC  (between  27°N  and  59°N;  117 
specimens):  Institut  fur  Seefischerei,  Hamburg  (ISH):  79/73 
(2).  MCZ:  53949  (1);  38282  (3);  38073  (3);  37633  (1);  37619  (2); 
37595  (1);  37520  (1);  37585  (2).  University  of  Maine,  Darling 
Center  (UMDC):  313-1  (1);  306-2  (1).  USNM:  21837-8  (2  syn- 
types of  Haloporphyrus  viola);  uncat.  (35);  31725  (1).  x-ray 
counts  only:  143250  (1);  45872  (1);  45845  (1);  45808  (1);  36163 
(1);  38142  (1);  38068  (1);  38064  (3);  38019  (1);  35595  (1);  35566 
(1);  33446  (4);  33443  (7);  33340  (3);  33337  (5);  33014  (1);  31768 
(1);  28612  (1);  28611  (1);  28610  (1);  28609  (1);  28608  (1);  24746 
(1).  Virginia  Institute  of  Marine  Sciences  (VIMS):  3458  (2); 
3243  (1);  1471  (7);  1460  (3);  872  (4);  870  (1);  uncat.  (3). 

EASTERN  NORTH  ATLANTIC  (between  50°N  and  60°N;  6 
specimens):  ISH:  112/74  (1);  111/74  (1);  146/74  (2);  745/74  (2). 

SOUTHEASTERN  PACIFIC  (between  0°S  to  56°S  and  70°W  to 
91°W;  123  specimens):  USNM:  uncat.  (120).  MCZ:  28610  (1) 
and  28611  (2)  (3  syntypes  of  Antimora  rhino). 

SOUTHERN  OCEANS  (specimens  from  southern  Atlantic,  In- 
dian, and  Pacific  oceans,  excluding  southeastern  Pacific  spec- 
imens; 101  specimens):  ISH:  2191/68  (2);  1250/66  (1);  1241/66 
(2);  1142/66  (11);  1129/66  (1);  361/71  (2);  344/71  (1);  286/71  (1); 


FIGURE  2.  Medial  view  of  first  gill  arch  of  left  side  to  il- 
lustrate location  of  gill  filament  measurement.  (A)  Antimora 
microlepis,  UW  17180,  off  Columbia  River,  46°N,  124°W,  310 
mm  SL;  (B)  Antimora  rostrata,  LACM  10985-5,  southwest  of 
New  Zealand,  56°19'S,  158°29'E,  330  mm  SL.  Drawn  by  Kei- 
ko  Hiratsuka  Moore. 


152/67  (8);  151/67  (3);  150/67  (1);  33/76  (5);  WH32/76  (5).  Los 
Angeles  County  Museum  (LACM):  10033  (12);  10032  (8); 
10985-5  (3);  uncat.  (9).  University  of  Florida  (UF-TABL):  503 
(1).  UMDC:  uncat.  station  numbers  01343  (1);  01342  (1);  00198 
(1);  00165  (1);  00157  (1);  00152  (1).  USNM:  188827  (1  syntype 
of  Antimora  australis);  188829  (3);  188823  (5);  188822  (2);  un- 
cat. (3).  British  Museum  Natural  History  (BMNH):  1887.12.- 
17.36  (holotype  of  Haloporphyrus  rostratus,  x-ray  only). 


RESULTS 

Antimora  may  be  divided  into  two  distinct 
species  instead  of  the  six  described,  based  on 
the  number  of  gill  filaments  and  secondarily  on 
the  ratio  of  gill  filament  length  to  standard 
length.  Other  measurements  can  be  used  but 
with  less  distinct  separation  of  species. 

COUNTS. — North  Pacific  specimens  possess 
90-103  gill  filaments  on  the  first  gill  arch,  com- 
pared with  76-90  in  specimens  from  the  other 


344 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  13 


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SMALL:  FISH  GENUS  ANTIMORA 


345 


•as) 


100 


200 


400 


500 


300 
Standard  length  (mm) 

FIGURE  3.  Regression  of  gill  filament  length  on  standard  length  in  Antimora  from  four  divisions  of  the  world  oceans:  North 
Pacific,  squares;  southeast  Pacific,  Southern  Ocean,  and  North  Atlantic,  dots.  North  Pacific:  y  =  0.03*  -  0.10,  n  =  51.  All 
other  oceans  (areas):  y  =  0.02*  -  0.70,  n  =  230. 


regions  (Table  1).  Number  of  gill  filaments  ap- 
pears to  be  constant  over  the  size  range  of  spec- 
imens in  all  populations. 

Only  slight  differences  between  samples  were 
found  from  the  four  geographical  areas  in  total 
vertebral  number,  number  of  gill  rakers  on  the 
first  arch,  and  anal  and  dorsal  fin  rays  (Table  2). 

Although  the  size  and  number  of  scales  have 
been  used  in  several  of  the  species  descriptions 
previously  mentioned,  they  are  not  useful  char- 
acters for  the  differentiation  of  these  species. 


Specimens  of  Antimora  are  very  fragile  and  on 
capture  the  scales  and  scale  pockets  do  not  re- 
main intact  for  use  as  a  reliable  character. 

MEASUREMENTS. — Specimens  collected  from 
the  eastern  and  western  North  Pacific  north  of 
latitude  10°N  have  a  gill  filament  length  relative- 
ly greater  than  that  in  fish  caught  elsewhere. 
Above  the  size  range  of  approximately  150  mm 
standard  length,  the  length  of  the  filaments  dis- 
tinctly separate  North  Pacific  fishes  from  all  oth- 
er groups.  Least  square  regression  lines  were  fit 


TABLE  2.     SUMMARY  OF  SELECTED  COUNTS  AND  LENGTH  PROPORTIONS  IN  Antimora  FROM  THE  FOUR  GEOGRAPHICAL 
REGIONS  SHOWN  IN  FIGURE  1  (lengths  presented  as  ratio  of  standard  length  to  size  of  part). 


N  Pacific 


SE  Pacific 


N  Atlantic 


S  Ocean 


Character 

X 

n 

SD 

X 

n 

SD 

X 

n 

SD 

X 

n          ! 

5D 

Snout  length 

11.9 

71 

0.86 

11.8 

105 

1.33 

12.7 

78 

1.16 

12.6 

93 

.53 

Predorsal  length 

3.9 

69 

1.47 

3.7 

108 

0.20 

3.7 

76 

0.17 

3.9 

94         ( 

).20 

Maxillary  length 

7.1 

70 

0.36 

6.9 

110 

0.50 

7.2 

76 

0.38 

7.4 

94         ( 

).49 

First  dorsal  fin  ray  length 

5.9 

47 

1.43 

7.1 

94 

1.58 

5.1 

63 

1.45 

6.1 

54 

.40 

Eye  diameter 

15.0 

66 

1.20 

15.3 

% 

1.34 

16.0 

69 

1.42 

16.2 

49 

.36 

Interorbital  width 

17.6 

38 

1.45 

18.6 

71 

1.61 

15.5 

32 

1.56 

18.3 

32 

.64 

Longest  gill  raker  length 

73.4 

23 

14.07 

- 

0 

- 

76.9 

1 

- 

103.0 

11        1 

3.00 

Total  number  of  vertebrae 

59.1 

47 

0.86 

58.8 

76 

0.96 

59.8 

64 

1.26 

59.6 

31 

.02 

Total  number  of  gill  rakers 

16.5 

38 

1.93 

16.2 

74 

1.29 

16.6 

46 

2.11 

16.0 

79 

.54 

Total  number  of  anal  fin  rays 

40.6 

41 

1.40 

39.3 

56 

1.39 

41.9 

67 

1.56 

40.0 

28 

.70 

Total  number  of  dorsal  fin  rays 

52.4 

40 

1.15 

51.7 

59 

1.37 

53.8 

60 

1.45 

53.2 

32 

.45 

346 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  13 


140 
13U 
120 


•  •       All  other  oceans  (areas) 


200  300  400  500 

Standard  Length  (mm) 

FIGURE  4.  Regression  of  head  length  on  standard  length  in  Antimora  from  four  divisions  of  the  world  oceans:  North  Pacific, 
squares;  North  Atlantic,  southeast  Pacific,  and  Southern  Ocean,  dots.  North  Pacific:  y  =  0.26x  +  2.22,  n  =  98.  All  other 
oceans  (areas):  y  =  0.23*  +  2.9432,  n  =  277. 


to  the  data,  and  the  regression  of  gill  filament 
length  on  standard  length  was  determined  fol- 
lowing the  methods  of  Zar  (1974)  (Fig.  3). 

Head  length  against  standard  length  (Fig.  4) 
also  separates  North  Pacific  specimens  from  all 
other  Antimora.  This  difference  is  most  marked 
in  fish  larger  than  200  mm  standard  length. 

Bean  (1890),  Carman  (1899),  Barnard  (1925), 
and  Pequeno  (1970)  noted  head  length  to  total 
length  or  standard  length  proportions  in  original 
descriptions  with  no  apparent  differences  among 
the  described  species. 

Comparisons  of  the  length  of  snout,  predorsal 
distance,  maxillary,  first  dorsal  fin  ray,  eye  di- 
ameter, longest  gill  raker,  and  width  of  interor- 
bital  are  summarized  in  Table  2.  None  of  the 
measurements  serve  unequivocally  to  separate 
the  North  Pacific  specimens  from  other  Anti- 
mora. Fish  from  all  other  areas  do  show  some 
differences:  southeast  Pacific,  North  Atlantic, 
and  Southern  Ocean  populations  may  have 
slightly  different  lengths  of  first  dorsal  fin  ray; 
North  and  southeast  Pacific  fish  appear  to  have 


larger  eyes  than  do  other  Antimora ;  and  North 
Atlantic  specimens  have  larger  interorbital 
widths.  Gill  raker  length  may  also  show  some 
differences,  however,  data  for  this  character  are 
incomplete. 

Within  the  range  of  A.  rostrata,  there  are  local 
differences  in  some  morphometric  characters, 
but  these  differences  are  not  consistent  through- 
out the  range  of  size  or  geography.  For  example, 
the  first  dorsal  fin  ray  to  standard  length  ratio  is 
higher  in  North  Atlantic  specimens  than  those 
from  other  areas.  This  longer  fin  ray  is  most  pro- 
nounced in  specimens  in  the  200^00  mm  range. 
North  Atlantic  fish  also  have  a  shorter  snout  and 
wider  interorbital  distance  over  certain  seg- 
ments of  their  size  range  as  compared  to  speci- 
mens from  other  regions.  Although  only  one 
specimen  from  low  latitudes  in  the  mid-Atlantic 
was  examined  in  this  study,  there  may  be  con- 
tact between  North  Atlantic  and  South  Atlantic 
populations  of  Antimora,  as  specimens  have 
been  taken  in  the  Bahamas  and  the  Gulf  of 
Guinea  (personal  communication,  Daniel  M. 


SMALL:  FISH  GENUS  ANTIMORA 


347 


B 


FIGURE  5.     (A)  Antimora  rostrata,  USNM  218479,  SL  346  mm,  male,  western  North  Atlantic,  36°39'N,  74°28'W,  1530-1610 
m;  (B)  Antimora  microlepis  CAS  32308,  SL  371  mm,  male,  off  California.  Drawn  by  Keiko  Hiratsuka  Moore. 


Cohen,  Systematics  Laboratory,  National  Ma- 
rine Fisheries  Service,  NOAA,  Washington, 
D.C.). 

CONCLUSIONS 
The  species  can  be  characterized  as  follows: 

Antimora  microlepis  Bean,  1890 

(Figure  55) 

Antimora  microlepis  BEAN,  1890:38  (type-locality:  51°23'N, 
130°34'W,  ALBATROSS  sta.  2860,  off  Cape  St.  James,  Queen 
Charlotte  Islands,  876  fms  [1602  m],  13  Aug.  1888). 

CHARACTERS. — Gill  filaments  on  first  gill  arch 
90-103;  gill  filaments  relatively  long,  regression 
equation  of  gill  filament  length  on  standard 
length  v  =  0.03*  -  0.10;  head  length  relatively 


long,  regression  equation  of  head  length  on  stan- 
dard length  y  =  0.26jc  +  2.22. 

RANGE. — Eastern  and  western  North  Pacific 
Ocean,  north  of  latitude  10°N. 

Antimora  rostrata  (Gunther,  1878) 

(Figure  5 A) 

Haloporphyrus  rostratus  GUNTHER,  1878:18  (type-locality: 
"midway  between  the  Cape  of  Good  Hope  and  Kerguelen 
Island;  east  of  the  mouth  of  the  Rio  de  la  Plata,"  CHAL- 
LENGER sta.  146,  1375  fms  [2515  m],  and  sta.  320,  600  fms 
[1097  m]). 

Haloporphyrus  viola  GOODE  AND  BEAN,  1878:257-260  (type- 
locality:  "outer  edge  of  Le  Have  Bank,  at  a  depth  of  four 
or  five  hundred  fathoms"  [approximately  43°N,  64°W]). 

Antimora  rhina  GARMAN,  1899:185-186  (type-locality:  Gulf  of 
Panama,  ALBATROSS  sta.  3353,  7°06'15"N,  80°34'W,  695  fms 
[1271  m],  sta.  3393,  7°15'N,  79°36'W,  1020  fms  [1865  m]). 


348 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  13 


Antimora  australis  BARNARD,  1925:499  (type-locality:  "off 
Cape  Point,  475-900  fathoms"  [869-1646  m]). 

Antimora  meadi  PEQUENO,  1970:14-16  (type-locality:  ANTON 
BRUUN  cruise  13,  between  34°06'S,  72°26'W,  and  34°12'S, 
72°25'W,  in  1400-1475  m,  3  Feb.  1966). 

CHARACTERS. — Gill  filaments  on  first  gill  arch 
76-90;  gill  filaments  relatively  short,  regression 
equation  of  gill  filament  length  on  standard 
length  y  =  0.02*  -  0.70;  head  length  relatively 
short,  regression  equation  of  head  length  on 
standard  length  y  =  0.23*  +  2.9432. 

RANGE. — All  oceans  except  the  North  Pacific 
north  of  10°N. 

ACKNOWLEDGMENTS 

I  am  thankful  for  the  advice  and  support  given 
me  by  Daniel  M.  Cohen  of  the  Systematics  Lab- 
oratory, National  Marine  Fisheries  Service 
(NMFS),  for  initially  suggesting  this  study,  and 
his  valuable  guidance  throughout  the  course  of 
this  work.  Tomio  Iwamoto  (CAS)  recommended 
that  I  examine  gill  filament  length,  arranged  for 
the  loan  of  specimens  from  the  California  Acad- 
emy of  Sciences,  and  reviewed  the  manuscript. 
Bruce  B.  Collette  (Systematists  Laboratory, 
NMFS)  and  Hugh  H.  DeWitt  (University  of 
Maine,  Orono)  also  reviewed  the  manuscript 
and  made  valuable  comments. 

Special  thanks  are  due  Charles  A.  Wenner  and 
John  A.  Musick  for  allowing  me  to  read  their 
then-unpublished  manuscript  on  the  life  history 
of  North  Atlantic  Antimora  and  examine  spec- 
imens in  their  collection  at  the  Virginia  Institute 
of  Marine  Science.  The  Allan  Hancock  Foun- 
dation and  Los  Angeles  County  Museum  pro- 
vided Antarctic  material,  William  Fink  (MCZ) 
provided  important  materials  from  the  North 
Atlantic  as  did  Alfred  Post  (ISH).  Richard  Ro- 
senblatt (SIO),  Arthur  Welander  (UW),  Carter 
Gilbert  (UF),  Jean  Dunn  (NMFS,  Seattle  Lab- 
oratory), and  the  National  Museum  of  Natural 
History  were  all  instrumental  in  providing  ad- 
ditional materials.  Keiko  H.  Moore  prepared  the 
illustrations  and  figures. 


LITERATURE  CITED 

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fishes  from  South  Africa.  Ann.  Mag.  Nat.  Hist.  ser.  9, 
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BEAN,  T.  H.  1890.  Scientific  results  of  explorations  by  the 
U.S.  Fish  Commission  Steamer  Albatross.  No.  XI — New 
fishes  collected  off  the  coast  of  Alaska  and  adjacent  region 
southward.  Proc.  U.S.  Natl.  Mus.  13:37^*5. 

COHEN,  D.  M.  1977.  Swimming  performance  of  the  gadoid 
fish  Antimora  rostrata  at  2400  meters.  Deep-Sea  Res. 
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GARMAN,  S.  1899.  Reports  on  an  exploration  off  the  west 
coasts  of  Mexico,  Central  and  South  America,  and  off  the 
Galapagos  Islands,  in  charge  of  Alexander  Agassiz,  by  the 
U.S.  Fish  Commission  Steamer  "Albatross"  during  1891, 
Lieut-Commander  Z.  L.  Tanner,  U.S.N.  Commanding.  Part 
26,  The  Fishes.  Mem.  Mus.  Comp.  Zool.  Harvard  24: 1—431. 

GOODE,  G.  B.,  AND  T.  H.  BEAN.  1878.  Descriptions  of  two 
gadoid  fishes,  Phycis  chest eri  and  Haloporphyrus  viola 
from  the  deep-sea  fauna  of  the  northwestern  Atlantic.  Proc. 
U.S.  Natl.  Mus.  1:256-260. 

GREY,  M.  1956.  The  distribution  of  fishes  found  below  a 
depth  of  2000  meters.  Fieldiana  Zool.  36:74-336. 

GUNTHER,  A.  1878.  Preliminary  notices  of  deep  sea  fishes 
collected  during  the  voyage  of  H.M.S.  Challenger.  Ann. 
Mag.  Nat.  Hist.  ser.  5,  2:17-28. 

HUBBS,  C.  L.,  AND  K.  F.  LAGLER.  1970.  Fishes  of  the  Great 
Lakes  region.  4th  ed.  Ann  Arbor:  Univ.  Michigan  Press. 
213  pp. 

IWAMOTO,  T.  1975.  The  abyssal  fish  Antimora  rostrata  (Giin- 
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PEQUENO-R.,  G.  1970.  Antimora  meadi  n.sp.  en  Chile.  Not. 
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SCHROEDER,  W.  C.  1940.  Some  deep  sea  fishes  from  the 
North  Atlantic.  Copeia  1940:231-238. 

SEDBERRY,  G.  R.,  AND  J.  A.  MUSICK.  1978.  Feeding  strate- 
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SVETOVIDOV,  A.  N.  1948.  Treskoobraznye.  Faune  SSSR, 
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OTS  63-1 1071.) 

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morid  fish,  Antimora  rostrata,  in  the  western  North  Atlan- 
tic. J.  Fish.  Res.  Bd.  Canada  34:2362-2368. 

ZAR,  J.  H.  1974.  Biostatistical  analysis.  Englewood  Cliffs, 
New  Jersey:  Prentice  Hall  Inc.  620  pp. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  14,  pp.  349-377;  18  figs.,  2  tables 


October  26,  1981 


THE  KAIANUS-GROUP  OF  THE  GENUS  CALLIONYMUS 

(PISCES:  CALLIONYMIDAE),  WITH  DESCRIPTIONS 

OF  SIX  NEW  SPECIES 

By 
Ronald  Fricke 

Saalestrasse  3  a,  D-3300  Braunschweig, 
Federal  Republic  of  Germany 


ABSTRACT:  The  kaianus -group  of  the  genus  Callionymus,  subgenus  Callionymus,  is  revised.  The  recognized 
species  and  their  ranges  are:  Callionymus  kaianus  Giinther,  1880  (Kai  Islands,  eastern  Indonesia);  Callionymus 
moretonensis  Johnson,  1971  (northern  half  of  Australia,  New  Ireland,  New  Caledonia);  Callionymus  whiteheadi 
n.sp.  (southwestern  Indonesia);  Callionymus  guentheri  n.sp.  (Philippine  Islands);  Callionymus  formosanus  n.sp. 
(Formosa  Strait,  Taiwan);  Callionymus  sokonumeri  Kamohara,  1936  (southern  Japan);  Callionymus  altipinnis 
n.sp.  (South  China  Sea:  China  coast);  Callionymus  ochiaii  n.sp.  (southern  Japan);  Callionymus  regani  Nakabo, 
1979  (Saya  de  Malha  Bank,  western-central  Indian  Ocean);  Callionymus  kotthausi  nom.  nov.  (for  Callionymus 
indicus  (Kotthaus,  1977),  a  secondary  homonym;  India);  Callionymus  africanus  (Kotthaus,  1977)  (east  Africa); 
Callionymus  bentuviai  n.sp.  (southern  Red  Sea);  Callionymus  carebares  Alcock,  1890  (northern  Indian  Ocean). 


INTRODUCTION 

The  dragonets  of  the  family  Callionymidae  are 
a  group  of  benthic  marine  fishes  (except  for  two 
euryhaline  species  which  enter  and  even  live  in 
freshwater  rivers).  About  130  species  are 
known.  The  two  largest  genera,  Callionymus 
and  Synchiropus,  are  nearly  circumtropical  in 
warm  and  temperate  seas,  but  some  species  also 
live  in  cold  waters;  e.g.,  Callionymus  lyra  and 
C.  maculatus  of  the  northern  Atlantic  follow  the 
warm  Gulf  Stream  to  Iceland  and  northern  Nor- 
way. Callionymids  usually  live  on  sand  or  mud 
bottoms,  sometimes  also  on  coral  sand  bottom 
in  coral  reefs,  or  among  seaweed,  from  very 
shallow  waters  and  even  tide  pools  down  to 
about  800  m. 

Callionymus  is  the  largest  genus  of  the  family, 
comprising  about  75  species.  Fricke  (1980)  dis- 
tinguished three  subgenera  (Callionymus,  Cal- 


liurichthys,  and  Spinicapitichthys)  which  differ 
principally  in  the  shape  of  the  preopercular 
spine.  The  subgenus  Callionymus  is  the  largest, 
comprising  about  55  species  which  can  be  ar- 
ranged into  various  species  groups.  The  kaia- 
nus-group  is  one  of  the  larger  species  groups  and 
contains  deepwater  mud-bottom  species  of  the 
Indian  and  western  Pacific  oceans. 

Seven  of  the  species  included  in  the  kaianus- 
group  have  been  described.  Callionymus  kaia- 
nus Giinther,  1880,  was  originally  described  from 
Kai  Islands,  west  of  New  Guinea,  and  later  re- 
corded from  India,  Indonesia,  Saya  de  Malha 
Bank  (western-central  Indian  Ocean),  Zanzibar, 
Arabian  Sea,  Japan,  Pescadore  Islands  (near 
Taiwan),  Gulf  of  Tonkin,  and  the  coast  of  China. 
Johnson  (1971)  described  a  new  subspecies, 
Callionymus  kaianus  moretonensis,  from  north- 
eastern Australia.  Callionymus  carebares  Al- 


[349] 


350 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


cock,  1890,  was  described  from  the  Bay  of  Ben- 
gal; C.  sokonumeri  Kamohara,  1936,  from 
Japan;  and  C.  regani  Nakabo,  1979,  from  the 
Say  a  de  Malha  Bank.  Under  the  generic  name 
Diplogrammus,  Kotthaus  (1977)  described  two 
further  species,  D.  africanus  and  D.  indicus. 

I  found  several  closely  allied  new  species, 
which  are  described  and  compared  in  the  pres- 
ent paper,  from  examination  of  specimens  (most 
previously  identified  as  Callionymus  kaianus)  in 
the  Australian  Museum,  Sydney  (AMS);  British 
Museum  (Natural  History),  London  (BMNH); 
California  Academy  of  Sciences,  San  Francisco 
(CAS);  Faculty  of  Agriculture  of  Kyoto  Univer- 
sity, Kyoto  (FAKU);  Fish  Collection,  Hebrew 
University  of  Jerusalem  (HUJF);  Institut  Royal 
des  Sciences  Naturelles,  Brussels  (IRSN);  Ma- 
rine Science  Laboratory,  Chinese  University  of 
Hong  Kong  (MSL);  Staatliches  Naturhisto- 
risches  Museum,  Braunschweig  (NMB);  Zoo- 
logisches  Institut  und  Zoologisches  Museum  der 
Universitat  Hamburg  (ZIM);  and  Museum  fur 
Naturkunde,  Zoologisches  Museum,  East  Berlin 
(ZMB). 

METHODS 

Methods  used  are  the  same  as  those  in  my 
previous  papers  (especially  Fricke  1980;  Fricke 
1981a). 

The  preopercular  spine  formula,  explained  by 
Fricke  (198  la),  is  calculated  by  the  following 
formula: 


where  a  is  the  number  of  antrorse  spines  at  the 
base,  b  is  the  number  of  points  or  serrae  at  the 
dorsal  edge,  c  is  the  number  of  points  or  serrae 
at  the  ventral  edge,  and  d  is  1  and  reflects  the 
main  tip  of  the  spine.  The  formula  treats  sim- 
plified left  spines.  Right  spines  have  to  be  treat- 
ed as  left  (e.g.,  the  number  of  antrorse  spines  at 
the  base,  a,  is  always  on  the  left  side  of  the 
formula). 

The  pectoral  fin  base  is  divided  in  two  by  a 
membrane  connecting  it  with  the  fifth  pelvic  fin 
ray.  The  formula  a/b  is  used  where  a  is  the  pec- 
toral fin  base  length  above  the  connecting  mem- 
brane, b  is  the  corresponding  length  below. 

THE  CALLIONYMUS  KAIANUS-GROUP 

The  Callionymus  kaianus  -group,  including 
the  new  species  described  in  this  paper,  com- 


prises thirteen  deepwater  mud-bottom  species 
in  the  subgenus  Callionymus  (see  Fricke 
1980:59)  distributed  in  the  Indian  and  west  Pa- 
cific oceans  (Figs.  1  &  2):  Callionymus  kaianus, 
C.  moretonensis,  C.  whiteheadi,  C.  guentheri, 
C.  sokonumeri,  C.formosanus,  C.  altipinnis,  C. 
ochiaii,  C.  regani,  C.  kotthausi,  C.  africanus, 
C.  bentuviai,  and  C.  carebares.  The  group  is 
characterized  by  the  presence  in  its  members  of 
large  eyes;  dorsal  and  anal  fin  formulae  D  IV  + 
viiij,  A  (viij-)viiij;  one  or  two  unbranched 
median  caudal  fin  rays  which  are  often  filamen- 
tous; a  characteristic  shape  of  the  preopercular 
spine  (see  Fig.  3);  and  (usually)  a  characteristic 
black  blotch  on  the  third  membrane  of  the  first 
dorsal  fin. 

Species  of  the  kaianus  -group  are  similar  to 
the  deepwater  mud-bottom  species  groups  of  the 
genus  Synchiropus  (e.g.,  phaeton -group,  alti- 
velis -group),  agreeing  with  them  in  some  aspects 
of  body  shape  and  even  color  markings  (black 
spot  on  third  membrane  of  first  dorsal  fin,  etc.). 
The  Synchiropus  species  groups  are  easily  dis- 
tinguished from  the  kaianus  -group  by  the  shape 
of  the  preopercular  spine  and  by  generic  differ- 
ences between  Callionymus  and  Synchiropus 
discussed  in  detail  in  my  revision  of  the  genus 
Synchiropus  (Fricke  1981b). 

Characters  and  distribution  of  the  species  of 
the  kaianus-group  are  compared  in  Tables  1-2. 
Further  distinguishing  features  (not  compared  in 
the  tables)  include  other  proportions,  the  pre- 
opercular spine  shape,  and  body  color  pattern. 

Key  to  the  Species  of  the 
Callionymus  kaianus-Group 

la.  Head  in  SL  2.7-3.1;  branchial  opening 
very  broad,  same  size  as  or  larger  than 
pupil;  upper  edge  of  preopercular  spine 
with  2  large  curved  points,  but  without 
a  small  antrorse  point  C.  carebares 

Ib.  Head  in  SL  3.4-4.6;  branchial  opening 
small,  about  Vs-Vz  of  pupil;  upper  edge 
of  preopercular  spine  with  1  small  an- 
trorse and  1  or  2  large  curved  points  __  2 

2a.  D2  and  anal  fins  very  high,  males  with 
convex  distal  margins  3 

2b.  D2  and  anal  fins  relatively  low,  with 
straight  distal  margins  6 

3a.  First  spine  of  D,  filamentous 4 

3b.  First  spine  of  D,  not  filamentous  ..  5 


FRICKE:   KA 1ANUS -GROUP  OF  GENUS  CALLIONYMUS 


351 


•  ochiaii        •carebares 
D  moretonensis 
A  kaianus 
v  af  ricanus 

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FIGURE  1.     Geographical  distribution  of  species  of  the  kaianus-group  of  the  genus  Callionymus. 


4a.  D2  relatively  low,  1st  ray  about  1.0,  5th 
ray  about  0.9  in  head  length;  preoper- 
cular  spine  with  2  curved  points  (and  1 
small  antrorse  point)  at  its  upper  side; 
black  blotch  on  3rd  membrane  of  D, 
large,  nearly  covering  entire  membrane; 
anal  fin  with  a  distal  black  streak  on  each 
membrane;  distal  %  of  anal  fin  dark 

brown,  tips  of  rays  white  

C.  formosanus 

4b.  D2  high,  1st  ray  about  1.1,  5th  ray  about 
0.8  in  head;  preopercular  spine  with  1 
large  curved  point  (and  1  small  antrorse 
point)  at  its  upper  side;  black  blotch  on 
3rd  membrane  of  D,  small,  distal  in  po- 
sition; anal  fin  without  a  distal  black 
streak  on  each  membrane;  distal  margin 
of  anal  fin  black,  tips  of  fin  rays  also 
black  C.  altipinnis 

5a.  Main  tip  of  preopercular  spine  long  and 
slender;  distal  half  of  anal  fin  black,  tips 
of  rays  white;  distal  margin  of  caudal  fin 
regular;  black  blotch  on  3rd  membrane 
of  D,  relatively  large,  central  in  position 

(not  reaching  distal  margin)  

C.  guentheri 


5b.  Main  tip  of  preopercular  spine  short; 
anal  fin  with  a  distal  black  streak  on  each 
membrane,  distal  %  of  membranes  and 
tips  of  fin  rays  brown;  distal  margin  of 
caudal  fin  irregular;  black  blotch  on  3rd 
membrane  of  D,  very  small,  extremely 
distal  in  position C.  sokonumeri 

6a.  Caudal  fin  convex,  without  filaments   __   7 
6b.  Caudal  fin  convex  or  slightly  pointed, 
with  1  or  2  filaments   10 

7a.  Anal  fin  with  a  small  distal  yellow  margin 
or  colorless;  sides  of  body  with  a  row  of 

large  indistinct  brownish  blotches 

C.  regani 

7b.  Anal  fin  with  a  broad  dark  brown  or 
black  margin;  sides  of  body  with  a  row 
of  small  distinct  black  blotches  8 

8a.  Main  tip  of  preopercular  spine  long  and 
slender;  lower  part  of  caudal  fin  colorless 
C.  kotthausi 

8b.  Main  tip  of  preopercular  spine  short; 
lower  part  of  caudal  fin  with  a  broad 
black  streak  9 

9a.  Pectoral  fin  base  with  a  large  dark  brown 


352 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol. No.  14 


Aregani         vguentheri 
T  kotthausi     A  altipinnis 
o  whiteheadi  •formosanus 
•  bentuviai 
D  sokonumeri 


FIGURE  2.     Geographical  distribution  of  species  of  the  kaianus-group  of  the  genus  Callionvmus. 


area;  back  marbled  with  olive-green; 
operculum  with  large  white  spots;  lower 
part  of  caudal  fin  black  distally;  D2  with 

a  basal  row  of  dark  blotches  

C.  whiteheadi 

9b.  Pectoral  fin  base  with  2  black  streaks; 
back  yellowish  brown,  with  white 
blotches  surrounded  by  semicircular 
dark  brown  bands;  operculum  with  small 
black  spots;  lower  part  of  caudal  fin  with 
a  broad,  curved  black  band,  but  color- 
less distally;  D2  without  a  basal  row  of 
black  spots  C.  africanus  (female) 

lOa.  First  spine  of  D,  with  a  long  filament    __ 

11 
lOb.  First  spine  of  D,  without  a  filament   __     13 

lla.  Main  tip  of  preopercular  spine  short, 
largest  point  on  its  dorsal  side  with  a  bas- 
al hook;  D2  colorless,  with  transverse 
white  lines;  sides  of  body  with  a  row  of 
distinct  black  blotches C.  kaianus 

lib.  Main  tip  of  preopercular  spine  long  and 
slender,  largest  point  on  its  dorsal  side 
without  a  basal  hook;  D,  with  rows  of 


blotches;  sides  of  body  with  a  row  of  in- 
distinct brownish  blotches    12 

12a.  Anal  fin  with  a  dark  margin;  D2  with  2 
rows  of  white  blotches  and  a  darkish  dis- 
tal margin;  main  tip  of  preopercular 
spine  about  2.5-3.0  times  as  long  as  larg- 
est point  at  its  dorsal  side;  body  with 
light  blotches  edged  with  semicircular 
dark  lines  C.  moretonensis 

12b.  Anal  fin  pale,  without  a  dark  margin;  D2 
with  a  basal  and  2-3  more  distal  rows  of 
dark  spots;  main  tip  of  preopercular 
spine  about  1.0-1.5  times  as  long  as  larg- 
est point  at  its  dorsal  side;  body  with 
minute  blackish  spots  forming  rings  and 
blotches  C.  ochiaii 

13a.  Second  membrane  of  D,  deeply  incised; 
caudal  fin  with  2  long  filaments  which  are 
nearly  twice  as  long  as  rest  of  fin;  D, 
with  2  or  more  black  blotches,  largest 
reaching  from  1st  to  4th  spine;  D2  with 
vertical  dark  streaks  C.  bentuviai 

13b.  Second  membrane  of  D,  not  incised; 
caudal  fin  with  1  or  2  relatively  short  fil- 


FRICKE:   KAIANUS-GROUP  OF  GENUS  CALLIONYMUS 


353 


aments  which  are  not  longer  than  rest  of 
fin;  D,  with  a  black  blotch  on  3rd  mem- 
brane (rarely  also  with  an  additional 
black  blotch  distally  on  the  same  mem- 
brane), occasionally  reaching  to  2nd 
spine;  D.,  with  rows  of  dark  and/or  light 
blotches  (but  without  vertical  dark 
streaks)  ..  14 

14a.  Lateral  line  in  area  behind  eye  with  a 
long  branch  running  downwards;  D,  with 
a  large  black  blotch  basally  on  3rd  mem- 
brane, 1  or  2  branches  of  which  reach 
2nd  membrane;  pectoral  fin  base  with  2 
dark  streaks;  anal  fin  with  a  distal  black 
margin  (usually  including  tips  of  fin 
rays);  caudal  fin  without  2  median  trans- 
verse black  lines;  D2  with  1  basal,  1  me- 
dian, and  1  distal  row  of  light  spots  and 

2  median  rows  of  black  spots 

C.  africanus  (male) 

14b.  Lateral  line  without  a  branch  in  the  post- 
orbital  area;  D,  with  a  relatively  small 
distal  black  blotch  on  3rd  membrane; 
pectoral  fin  base  with  a  dark  area;  distal 
half  of  anal  fin  black,  tips  of  fin  rays 
white;  caudal  fin  with  2  median  trans- 
verse black  lines;  D2  with  a  basal  and  a 
median  row  of  black  blotches __. 

C.  guentheri  (female) 

Callionymus  carebares  Alcock,  1890 

(Figures  4-5) 

Callionymus  carehares  ALCOCK,  1890:209  ("off  Madras  coast, 
98-102  fms"):  1898:73;  1899: pi.  20,  fig.  4:  REGAN  1906:329 
(Sea  of  Oman,  98-180  fms  [179-329  m]):  SMITH  1963:555, 
pi.  84K  (after  Alcock). 

MATERIAL  EXAMINED.— Syntypes:  BMNH  1890.11.28.18- 
24,  2  6 ,  5  9,  "Investigator,"  off  Ganjam  Coast,  India. 

Other  specimens:  BMNH  1903.5.14.34,  1  spec. ,-39.0  mm 
SL,  J.  W.  Townsend,  Karachi.  BMNH  1903.9.24.2-4,  3  spec.. 
J.  W.  Townsend,  Iranian  Mekran  coast,  Gulf  of  Oman 
(25°19'N,  58°21'E),  98  fms  (179  m).  BMNH  1904.5.25.218- 
220,  3  spec.,  J.  W.  Townsend,  Sea  of  Oman,  180  fms  (329  m). 
BMNH  1939.5.24.1384,  1  spec.,  John  Murray  Exped..  23  Nov. 
1933,  Gulf  of  Oman,  193  m.  BMNH  1939.5.24.1385-1409,  24 
spec.,  John  Murray  Exped.,  Arabian  Sea,  135-183  m.  BMNH 
1939.5.24.1410-1421,  15  spec.,  John  Murray  Exped.,  Gulf  of 
Aden,  220  m.  IRSN  1797,  2  9,  M.  Frank,  4  Apr.  1894,  Gulf 
of  Bengal.  FMNH  5740,  1  9,  J.  W.  Townsend,  1906,  Sea  of 
Oman  (Dr.  D.  J.  Stewart,  FMNH,  kindly  examined  the  spec- 
imen). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  very  large  branchial  opening  (same 
size  as,  or  larger  than,  pupil),  an  unusually  large 


^ — ' 


FIGURE  3.  Preopercular  spines  of  species  of  the  kaianus- 
group.  The  main  characteristics  are  the  straight  ventral  side 
of  the  spine,  the  strong  straight  antrorse  point,  the  upcurved 
main  tip,  and  the  antrorse  spine  in  combination  with  two  (rare- 
ly one)  large  curved  points  on  the  dorsal  side.  (A)  Left  pre- 
opercular  spine  of  Callionymus  kaianus.  (R)  Left  preopercu- 
lar  spine  of  C.  guentheri.  (C)  Left  preopercular  spine  of  C. 
africanus.  (D)  Left  preopercular  spine  of  C.  moretonensis . 
(E)  Left  preopercular  spine  of  the  most  primitive  species  of 
the  kaianus  -group,  C.  curehares  (without  an  antrorse  point 
on  the  dorsal  side). 


head  (head  length  in  SL  2.7-3.1),  and  two  large 
curved  points,  but  without  a  small  antrorse  spine 
at  dorsal  edge  of  preopercular  spine. 

DESCRIPTION. — Counts  and  measurements 
(see  also  Table  1):  D,  IV;  D2  viii.l;  A  viii,l;  P, 
ii-iji,14-16,iii;  P,  1,5;  C  ii,2-3,0-ii,2-4,iii. 

Body  elongate  and  depressed.  Head  very 
large,  depressed,  2.7-3.1  in  SL.  Eye  large,  2.7- 
3.5  in  head.  Pupil  large,  about  2.5  in  eye.  Bran- 
chial opening  very  large,  same  size  as  pupil  or 
larger.  Occipital  region  with  a  bony  plate  and 
two  low  bony  protuberances.  Preopercular  spine 
nearly  as  long  as  eye  diameter,  with  a  long, 
slightly  upcurved  main  tip,  two  curved  points  at 
its  dorsal  side  and  a  large  antrorse  spine  at  its 


354 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


S- 


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FRICKE:   KA1ANUS-GROUP  OF  GENUS  CALLIONYMUS 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


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FIGURE  4.  Callionymux  carebares,  IRSN  1797,  Gulf  of  Bengal.  Specimen  1,  female,  85.8  mm  SL:  (A)  lateral  view;  (B) 
dorsal  view;  (C)  ventral  view;  (D)  left  preopercular  spine;  (E)  right  preopercular  spine.  Specimen  2,  female,  86.2  mm  SL:  (F) 
right  preopercular  spine. 


base  (formula:   1- 


-1;  see  Fig.  4D-F).  Lat- 


eral line  reaching  from  hind  margin  of  eye  to 
base  of  caudal  fin;  the  line  of  the  opposite  side 
is  interconnected  by  a  transverse  branch  across 
the  occipital  region.  Caudal  peduncle  length  5.8- 
6.6  in  SL,  minimal  caudal  peduncle  depth  21.0- 
22.6  in  SL. 

First  spine  of  first  dorsal  fin  nearly  as  long  as 
first  ray  of  second  dorsal  fin,  filamentous  only 
in  females.  Distal  margin  of  second  dorsal  fin 
straight.  Anal  fin  beginning  on  vertical  through 
second  ray  of  second  dorsal  fin.  Distal  margin 
of  caudal  fin  slightly  convex,  the  two  median 
rays  elongate  but  usually  not  filamentous.  Outer 
edge  of  pelvic  fin  convex;  longest  pelvic  fin  ray 
reaching  to  base  of  first  anal  fin  ray.  Pectoral  fin 
reaching  nearly  to  fifth  ray  of  second  dorsal  fin 
when  laid  back. 


Color  in  alcohol.  Head  and  body  dark  gray, 
belly  white.  Eye  darkish.  First  dorsal  fin  in  male 
monochromatic  dark,  in  female  nearly  colorless, 
with  a  large  distal  dark  blotch  reaching  from  sec- 
ond to  fourth  spine.  Second  dorsal  fin  colorless, 
distal  margin  darkish.  Distal  one-third  of  caudal 
fin  dark.  Distal  two-thirds  of  anal  fin  black,  anal 
fin  base  colorless.  Pelvic  fin  colorless,  pectoral 
fin  with  few  dark  spots. 

DISTRIBUTION. — Northern  parts  of  Indian 
Ocean:  Gulf  of  Aden,  Gulf  of  Oman,  coast  of 
India,  Arabian  Sea  (see  Fig.  1);  135-330  m  on 
muddy  bottoms. 

DISCUSSION. — Callionymus  carebares  seems 
to  be  the  most  primitive  member  of  the  kaianus- 
group  based  on  the  shapes  of  its  preopercular 
spine  (no  antrorse  spine  at  its  dorsal  side)  and 
caudal  fin  (often  no  median  unbranched  ray 


FRICKE:   KAIANUS-GROUP  OF  GENUS  CALLIONYMUS 


357 


FIGURE  5.  First  dorsal  fin  in  Callionymus  carebares.  (A) 
BMNH  1890.11.28.18,  male,  95.0  mm  SL,  syntype,  Ganjam 
coast,  India.  (B)  BMNH  1890.11.28.19,  female,  86.2  mm  SL, 
syntype,  Ganjam  coast,  India. 


present).  It  is  also  unique,  however,  in  its  very 
large  head  and  its  extremely  large  branchial 
opening  (which  is  porelike  and  very  small  in  oth- 
er callionymid  fishes).  Therefore,  it  seems  to 
belong  to  another  evolutionary  branch  in  the 
kaianus -group,  and  I  assign  it  to  a  subgroup  of 
its  own.  Juvenile  specimens,  which  have  a 
smaller  head  and  a  smaller  branchial  opening, 
are  more  similar  to  the  other  species  of  the 
kaianus -group. 

Callionymus  kaianus  Giinther,  1880 

(Figure  6) 

Callionymus  kaianus  GUNTHER,  1880:44,  pi.  19,  fig.  B  (Kai 
Is.,   129  fms  [236  m]);  DE  BEAUFORT  1951:66-67,  fig.   12 


(after  Gunther);  SMITH  1963:553,  pi.  847  (in  part;  after  Giin- 
ther);  SUWARDJI  1965:308-310  (Kai  Is.,  180-290  m). 
MATERIAL  EXAMINED.— Holotype:   BMNH   1879.5.14.565, 
1  6,  128.6  mm  SL,  Challenger  Exped.,  Kai  Is. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening,  a  short 
head  (about  4.6  in  SL),  preopercular  spine  with 
a  small  antrorse  and  two  large  curved  points  on 
dorsal  side,  straight  distal  margin  of  second  dor- 
sal fin,  two  short  caudal  fin  filaments,  long  fila- 
mentous first  spine  of  first  dorsal  fin,  short  main 
tip  of  preopercular  spine,  and  pale  anal  fin. 

DESCRIPTION. — Counts  and  measurements 
(see  also  Table  2):  D,  IV;  D2  viii.l;  A  viii.l;  P, 
ii,17-18,ii;  P2  1,5;  C  ii,2,ii,3,iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, about  4.6  in  SL.  Eye  large,  2.35  in 
head.  Pupil  relatively  small,  3.3  in  eye.  Bran- 
chial opening  of  normal  size,  about  two  times  in 
pupil.  Occipital  region  with  two  low  bony  pro- 
tuberances. Preopercular  spine  1.45  in  eye  di- 
ameter, with  a  relatively  short,  slightly  upcurved 
main  tip,  a  small  antrorse  and  two  large  curved 
points  (the  larger  with  a  small  additional  basal 
point)  on  its  dorsal  side  and  a  large  antrorse 

spine  at  its  base  (formula:   1 1;  see  Fig. 

6B).  Lateral  line  reaching  from  area  behind  eye 
to  end  of  third  branched  caudal  fin  ray  (seen 


10mm 


B 


••^^•MMM 

2mm 


FIGURE  6.     Callionymus  kaianus,  BMNH  1879.5.14.565,  holotype,  male,  128.6  mm  SL,  Kai  Islands:  (A)  lateral  view;  (B) 
left  preopercular  spine. 


358 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


10  mm 


B 


10mm 


2mm 


D       2mm 


FIGURE  7.     Callionymus  moretonensis,  NMB  37074,  69.4  mm  SL,  New  Caledonia:  (A)  lateral  view;  (B)  dorsal  view,  (C)  left 
preopercular  spine:  (D)  right  preopercular  spine. 


from  above);  the  line  of  one  side  is  intercon- 
nected to  its  opposite  member  by  a  transverse 
branch  across  the  occipital  region  and  one 
across  the  dorsal  side  of  caudal  peduncle.  Cau- 
dal peduncle  length  5.2  in  SL,  minimal  caudal 
peduncle  depth  23.8  in  SL. 

First  spine  of  first  dorsal  fin  long  and  filamen- 
tous. Distal  margin  of  second  dorsal  fin  straight. 
Anal  fin  beginning  on  the  vertical  through  mid- 
base  of  second  membrane  of  second  dorsal  fin. 
Distal  margin  of  caudal  fin  pointed,  the  two  un- 
branched  median  rays  filamentous  (filaments  rel- 


atively short).  Distal  margin  of  pelvic  fin  con- 
vex; longest  pelvic  fin  ray  only  reaching  the  anal 
papilla.  Pectoral  fin  reaching  to  second  anal  fin 
ray  when  laid  back. 

Color  in  alcohol.  Head  and  body  light  brown, 
back  with  some  lighter  blotches.  Belly  white, 
thorax  brownish.  Eye  light  gray.  Pectoral  fin 
base  with  a  large  dark  blotch.  A  row  of  dark 
blotches  in  groups  along  body  side.  Operculum 
with  some  dark  spots,  head  with  indistinct 
brownish  blotches  and  lines.  First  dorsal  fin 
light,  with  a  black  blotch  on  third  membrane,  a 


FR1CKE:   KAIAN US-GROUP  OF  GENUS  CALLIONYMVS 


359 


basal  branch  of  which  reaches  nearly  to  second 
spine.  Second  dorsal  fin  colorless,  with  white 
lines  in  characteristic  arrangement  (see  Fig.  6A). 
Anal  fin  colorless,  with  few  yellow  pigment  on 
the  membranes  between  the  five  posterior  rays. 
Upper  and  median  parts  of  caudal  fin  with  dark 
spots  and  blotches;  lower  part  with  a  broad 
curved  dark  band.  Distal  part  of  pelvic  fin  dark; 
pectoral  fin  colorless. 

DISTRIBUTION. — Kai  Islands  (west  of  New 
Guinea),  at  three  different  localities  (Fig.  1); 
180-290  m,  muddy  bottom. 

DISCUSSION. — C.  kaianus  differs  from  C. 
moretonensis  of  northern  Australia,  the  species 
geographically  nearest,  by  its  shorter  main  tip 
of  preopercular  spine,  and  by  various  color 
markings  (e.g.,  pale  anal  fin  without  a  black  dis- 
tal margin);  it  differs  from  C.  whiteheadi  by  the 
presence  of  filaments  in  the  first  dorsal  and  cau- 
dal fins,  the  barbed  largest  point  on  dorsal  side 
of  preopercular  spine,  and  various  color  mark- 
ings. 

Specimens  referred  to  C.  kaianus  from  the 
Gulf  of  Thailand  seem  to  belong  to  another 
species. 

Callionymus  moretonensis  Johnson,  1971 

(Figure  7) 

Callionymus  calauropomus:  (nee  Richardson,  1844)  PETERS, 
1876:841  (New  Ireland);  JORDAN  AND  SEALE  1905:415  (after 
Peters);  FOWLER  1928:422  (after  Peters);  MUNRO  1958:253 
(after  Peters). 

Callionymus  kaianus  moretonensis  JOHNSON,  1971: 108-J  13, 
figs.  1-2  (s  Queensland);  1973:217-230  (biology). 

MATERIAL  EXAMINED.—  Holotype:  AMS  115608-001,  1 
spec.,  158.3  mm  SL,  C.  R.  Johnson,  1  Aug.  1969,  E  of  Cape 
Moreton,  Queensland,  68-72  fms  (124-132  m).  Paratypes: 
CAS  24764,  1  spec.,  131.2  mm  SL;  CAS  24765,  1  spec.,  134.0 
mm  SL;  CAS  24766,  1  spec.,  141.7  mm  SL:  CAS  24767,  1 
spec.,  164.9  mm  SL;  all  with  same  data  as  holotype. 

Other  specimens:  BMNH  1892.1.14.26-27,  2  spec.,  52.4- 
58.2  mm  SL,  Mr.  Walker,  Holothuria  Banks  (NW  Australia). 
ZMB  9399,  1  d,  1  juv.,  35.0-81.9  mm  SL,  R/V  GAZELLE, 
"shortly  before  the  year  1876,"  New  Ireland.  NMB  37074,  1 
spec.,  69.4  mm  SL,  P.  Fourmanoir,  1979,  Havannah,  s  New 
Caledonia,  150  m. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.4-4.7  in  SL);  preopercular  spine  with  a  small 
antrorse  and  two  large  curved  points  on  its  dor- 
sal side  and  a  long,  slender  main  tip;  second 
dorsal  fin  with  a  nearly  straight  distal  margin; 
caudal  fin  with  two  short  median  filaments;  first 
dorsal  fin  with  a  relatively  long,  filamentous  first 
spine;  anal  fin  with  a  dark  distal  margin;  and 


second  dorsal  fin  with  rows  of  white  blotches 
and  a  dark  distal  margin. 

DESCRIPTION. — Counts  and  measurements 
(see  also  Table  2):  D,  IV;  D2  viii,l;  A  vii,l-viii,l; 
P,  ii,  18-20;  P2  1,5;  C  i-ii,3-4,  (0-)i-ii,2-3,ii-iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, about  3.4-4.7  in  SL.  Eye  large,  2.0-2.4 
in  head.  Pupil  relatively  small,  2.9-3.0  in  eye. 
Branchial  opening  of  normal  size,  about  2-3 
times  in  pupil.  Occipital  region  with  two  low 
bony  protuberances.  Preopercular  spine  about 
1.1-1.2  in  eye  diameter,  with  a  long  slightly  up- 
curved  main  tip,  a  small  antrorse  and  two  large 
curved  points  at  its  dorsal  side,  and  a  large  an- 
trorse spine  at  its  base  (formula:  1 1;  see 

Fig.  1C,  D).  Lateral  line  reaching  from  area  be- 
hind eye  to  end  of  third  branched  caudal  fin  ray 
(counted  from  above);  the  line  of  the  opposite 
side  is  interconnected  by  a  transverse  branch 
across  the  occipital  region  and  one  across  the 
dorsal  side  of  the  caudal  peduncle.  Caudal  pe- 
duncle length  5.8-6.5  in  SL,  minimal  caudal  pe- 
duncle depth  23.0-30.0  in  SL. 

First  spine  of  first  dorsal  fin  relatively  long 
and  filamentous  (in  adults).  Distal  margin  of  sec- 
ond dorsal  fin  straight.  Anal  fin  beginning  on  a 
vertical  through  second  ray  of  second  dorsal  fin. 
Distal  margin  of  caudal  fin  convex  in  small  spec- 
imens (somewhat  pointed  in  adults),  with  two 
short  median  filaments.  Distal  margin  of  pelvic 
fin  convex;  longest  pelvic  fin  ray  reaching  nearly 
to  midbase  of  first  membrane  of  anal  fin.  Pec- 
toral fin  reaching  to  fourth  ray  of  second  dorsal 
fin. 

Color  in  alcohol.  Head  and  body  light  brown, 
ventral  side  of  body  and  belly  whitish.  Thorax 
white.  Back  with  white  spots  bordered  by  semi- 
circular black  lines.  Eye  grayish.  A  row  of  ir- 
regular brownish  spots  along  sides  of  body. 
Head  with  small  white  spots.  Some  dark  spots 
at  upper  part  of  pectoral  fin  base.  First  dorsal 
fin  light  brown,  with  a  large  black  blotch  on  third 
membrane,  a  basal  branch  of  which  reaches  to 
second  spine,  and  some  smaller  white  blotches 
surrounding  it.  Second  dorsal  fin  with  two  or 
three  rows  of  white  spots  and  a  darkish  distal 
margin.  Anal  fin  colorless,  with  a  black  distal 
border.  Pelvic  fin  with  small  dark  spots  on  distal 
parts  of  fourth  and  fifth  rays.  Pectoral  fin  col- 
orless or  with  three  vertical  rows  of  darkish 
spots.  Caudal  fin  whitish,  with  some  dark  spots 
forming  a  broad  curved  band  in  the  upper  part, 
a  broad  curved  black  band  in  the  lower  part. 


360 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


DISTRIBUTION. — Northern  half  of  Australia 
(Holothuria  Banks  to  southern  Queensland), 
New  Caledonia,  and  New  Ireland  (see  Fig.  1); 
101-150  m,  mud  bottom. 

DISCUSSION. — The  differences  between  C. 
kaianus  and  C.  moretonensis  are  discussed  in 
the  description  of  the  former  species.  Callio- 
nymus  moretonensis  differs  from  C.  whiteheadi 
by  the  presence  of  filaments  in  first  dorsal  and 
caudal  fins,  by  the  shape  of  the  main  tip  of  the 
preopercular  spine,  and  by  color  markings. 

The  record  of  Synchiropus  calauropomus 
(Richardson,  1844)  from  New  Ireland  (Peters 
1876)  is  based  on  two  specimens  of  Callionymus 
moretonensis.  Synchiropus  calauropomus  does 
not  occur  in  that  area. 

Johnson  (1971)  originally  described  C.  more- 
tonensis as  a  subspecies  of  C.  kaianus,  but  the 
differences  are  sufficient  to  regard  C.  moreton- 
ensis as  a  distinct  species. 

Callionymus  whiteheadi  new  species 

(Figure  8) 

^Callionymus  kaianus:   (non  Giinther,    1880)   WEBER    1913: 
(Madura  Sea,  7°2.6'S,  115°23.6'E,  100  m). 

MATERIAL  EXAMINED.—  Holotype:  BMNH  1980.6.20.1, 
112.3  mm  SL,  P.  J.  P.  Whitehead,  14  July  1979,  off  Bali 
(8°50'S,  114°14'E),  1 10-220  m.  Paratype:  BMNH  1980.11.25.2, 
1  spec.,  105.5  mm  SL,  P.  J.  P.  Whitehead,  summer  1979,  SE 
coast  of  Java  (near  type-locality). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.9-4.2  in  SL);  preopercular  spine  with  a  small 
antrorse  and  one  or  two  large  curved  points,  and 
a  short  main  tip;  second  dorsal  fin  with  a  nearly 
straight  distal  margin;  caudal  fin  with  a  convex 
distal  margin,  but  without  filaments;  first  dorsal 
fin  without  a  filament  and  with  a  basal  black 
blotch  on  third  membrane;  light  brown  cheeks 
with  few  dark  spots;  anal  fin  with  a  black  distal 
margin;  second  dorsal  fin  with  a  basal  row  of 
dark  spots;  and  sides  of  body  with  a  row  of  dis- 
tinct black  blotches. 

DESCRIPTION. — Counts  and  measurements 
(see  also  Table  1):  D,  IV;  D2  viii.l;  A  viii,l;  P, 
i-ii,15-17,ii;  P2  1,5;  C  ii,3,i-ii,2-3,iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.9-4.2  in  SL.  Eye  large,  2.2-2.3  in 
head.  Pupil  relatively  small,  about  3.2  in  eye. 
Branchial  opening  of  normal  size,  about  3  times 
in  pupil.  Occipital  region  with  two  low  bony  pro- 
tuberances. Preopercular  spine  1.7-1.9  in  eye 
diameter,  with  a  short,  slightly  upcurved  main 


tip,  a  small  antrorse  and  one  or  two  large  curved 
points  at  its  dorsal  side,  and  a  large  antrorse 

spine  at  its  base  (formula:    1 — 1;  see  Fig. 

8#).  Lateral  line  reaching  from  area  behind  eye 
to  third  branched  caudal  fin  ray  (counted  from 
above);  the  line  of  the  opposite  side  is  intercon- 
nected by  a  transverse  branch  across  the  occip- 
ital region  and  another  across  the  dorsal  side  of 
caudal  peduncle.  Caudal  peduncle  length  5.5- 
6.1  in  SL,  minimal  caudal  peduncle  depth  25.5- 
26.3  in  SL. 

First  spine  of  first  dorsal  fin  somewhat  longer 
than  first  ray  of  second  dorsal  fin,  but  not  fila- 
mentous. Distal  margin  of  second  dorsal  fin 
nearly  straight.  Anal  fin  beginning  on  the  vertical 
through  second  ray  of  second  dorsal  fin.  Distal 
margin  of  caudal  fin  convex;  no  median  fila- 
ments. Distal  margin  of  pelvic  fin  convex;  long- 
est pelvic  ray  only  reaching  to  anal  papilla  when 
laid  back.  Pectoral  fin  reaching  to  midbase  of 
second  membrane  of  second  dorsal  fin  when  laid 
back. 

Color  in  alcohol.  Head  and  body  dark  olive- 
green;  lower  surface  of  body  white.  Back  mar- 
bled with  brown.  Thorax  and  belly  white.  A  dark 
area  at  upper  part  of  pectoral  fin  base.  Some 
large  whitish  spots  bordered  by  black  on  oper- 
culum.  Head  with  dark  brown  spots  and  lines. 
Eye  dorsally  black,  ventrally  dark  blue.  A  row 
of  irregular  black  blotches  along  sides  of  body. 
First  dorsal  fin  olive-green;  first  spine  marbled 
alternating  black  and  white;  a  large  black  blotch 
basally  on  third  membrane,  a  basal  branch  of 
which  reaches  to  second  spine.  Second  dorsal 
fin  with  two  rows  of  indistinct  darkish  blotches 
and  a  basal  row  of  distinct  blackish  blotches. 
Anal  fin  white,  with  a  broad  distal  black  margin. 
Caudal  fin  rays  in  upper  part  of  caudal  fin  with 
black  blotches;  lower  part  of  caudal  fin  blackish 
distally.  Distal  two-thirds  of  pelvic  fin  darkish; 
upper  half  of  pectoral  fin  with  four  vertical  rows 
of  black  spots. 

DISTRIBUTION. — Bali  and  southeastern  Java, 
possibly  also  Madura  Sea  (see  Fig.  2);  110-220 
m,  mud  bottom. 

DISCUSSION. — The  differences  between  C. 
whiteheadi,  C.  kaianus,  and  C.  moretonensis 
have  been  discussed  in  the  descriptions  of  the 
last  two  species.  C.  whiteheadi  differs  from  C. 
regani  by  the  shape  of  the  preopercular  spine 
and  by  various  color  markings  (e.g.,  black  bor- 
der of  anal  fin;  shape  and  position  of  black 
blotch  in  first  dorsal  fin;  color  patterns  of  second 


FRICKE:   KAIANUS-GROUP  OF  GENUS  CALL1ONYMUS 


361 


•^^^^^ 

10mm 


B 


2mm 


FIGURE  8.     Callionymus  whiteheadi,  holotype,  BMNH  1980.6.20.1,  112.3  mm  SL,  off  Bali:  (A)  lateral  view;  (B)  left  pre- 
opercular  spine. 


dorsal  and  caudal  fins;  body  color  pattern,  etc.). 
It  differs  from  C.  guentheri  in  lacking  caudal  fin 
filaments,  in  the  different  shape  of  the  preoper- 
cular  spine,  and  in  various  color  markings. 

ETYMOLOGY. — This  new  species  is  named  af- 
ter Dr.  Peter  J.  P.  Whitehead,  British  Museum 
(Natural  History),  who  collected  the  type-spec- 
imens and  allowed  me  to  examine  them. 

Callionymus  regani  Nakabo,  1979 

(Figure  9) 

Callionymus  kaianus:  (non  Giinther,  1880)  REGAN  1908:248 
(Saya  de  Malha  Bank,  123  fms  [225  m]);  SMITH  1963 
(part):553  (after  Regan). 

Callionymus  regani  NAKABO,  1979:231-234,  fig.  1,  table  1 
(Saya  de  Malha  Bank). 

MATERIAL  EXAMINED.— BMNH  1908.3.23.263,  1  d,  101.3 
mm  SL,  Gardiner-Expedition,  Saya  de  Malha  Bank  (western- 
central  Indian  Ocean),  "over  123  fms"  (225  m). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(about  4.1  in  SL);  preopercular  spine  with  a 
small  antrorse  and  two  large  curved  points  at 
dorsal  side,  and  a  short  main  tip;  second  dorsal 
fin  with  a  nearly  straight  distal  margin;  caudal 
fin  with  a  convex  distal  margin  but  without  fil- 


aments; first  dorsal  fin  without  a  filament  and 
with  a  distal  black  blotch  on  third  membrane 
surrounded  by  white  lines;  dark  brown  cheeks 
with  characteristic  light  blotches;  colorless  anal 
fin  without  a  distal  black  margin;  second  dorsal 
fin  with  rows  of  white  spots  and  lines;  and  sides 
of  body  with  few  large  indistinct  brownish 

blotches. 
DESCRIPTION. — Counts  and  proportions  (see 

Table  1):  D,  IV;  D2  viii.l;  A  viii.l;  P,  i-iii,17- 
19,0-i;  P2  1,5;  C  0-ii,3,i-ii,2-3,ii-iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 4.1  in  SL.  Eye  large,  about  2.7  in  head. 
Pupil  relatively  small,  3.4  in  eye  diameter.  Bran- 
chial opening  of  normal  size,  about  3  in  pupil. 
Occipital  region  with  two  low  bony  protuber- 
ances. Preopercular  spine  1.4  in  eye  diameter, 
with  a  short,  slightly  upcurved  main  tip,  a  small 
antrorse  and  two  larger  curved  points  at  its  dor- 
sal side,  and  a  large  antrorse  spine  at  its  base 
3 


(formula:    1- 


-1;  see  Fig.  9B).  Lateral  line 


reaching  from  area  behind  eye  to  end  of  third 
branched  caudal  fin  ray  (seen  from  above);  the 
line  of  the  opposite  side  is  interconnected  by  a 
transverse  branch  across  the  occipital  region 
and  another  across  the  caudal  peduncle.  Caudal 


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10mm 


B 


2mm 


FIGURE  9.     Callionymus  regani,  BMNH  1908.2.23.263,  male,  101.3  mm  SL,  Saya  de  Malha  Bank:  (A)  lateral  view;  (B)  left 
preopercular  spine. 


peduncle  length  6.25  in  SL;  minimal  depth  of 
caudal  peduncle  22.5  in  SL. 

First  spine  of  first  dorsal  fin  somewhat  longer 
than  first  ray  of  second  dorsal  fin,  but  not  fila- 
mentous. Distal  margin  of  second  dorsal  fin 
nearly  straight.  Anal  fin  beginning  on  vertical 
through  second  ray  of  second  dorsal  fin.  Distal 
margin  of  caudal  fin  nearly  convex;  no  median 
filaments.  Distal  margin  of  pelvic  fin  convex; 
longest  pelvic  fin  ray  reaching  to  base  of  first 
anal  fin  ray  when  laid  back.  Pectoral  fin  reaching 
to  midbase  of  third  membrane  of  second  dorsal 
fin  when  laid  back. 

Color  in  alcohol.  Head  and  body  brown.  Belly 
white,  thorax  light  brown.  Three  to  four  large 
indistinct  darkish  areas  along  sides  of  body. 
Head  brown;  males  with  many  light  blotches, 
females  with  few  light  blotches.  Occipital  region 
with  dark  spots.  Eye  posteriorly  gray,  anteriorly 
yellowish.  A  large  dark  brown  blotch  at  pectoral 
fin  base.  First  dorsal  fin  pale;  about  four  hori- 
zontal dark  lines  on  first  and  second  membranes. 


A  black  blotch  distally  on  third  membrane,  oc- 
casionally reaching  to  posterior  part  of  second 
membrane;  distal  edge  of  third  membrane  also 
black.  Remaining  parts  of  third  and  fourth  mem- 
branes covered  with  curved  white  lines.  Second 
dorsal  fin  mostly  colorless,  with  rows  of  white 
blotches  and/or  lines.  Anal  fin  pale;  distal  margin 
yellowish.  Caudal  fin  pale,  occasionally  with 
two  darkish  blotches  at  upper  edge;  lower  part 
sometimes  dusky.  Distal  parts  of  fourth  and  fifth 
rays  of  pelvic  fin  darkish;  pectoral  fin  colorless. 

DISTRIBUTION. — Saya  de  Malha  Bank,  west- 
ern-central Indian  Ocean  (see  Fig.  2);  126- 
225  m. 

DISCUSSION. — The  differences  from  Callio- 
nymus whiteheadi  were  discussed  in  the  descrip- 
tion of  that  species.  Callionymus  regani  differs 
from  C.  africanus  by  the  absence  of  caudal  fin 
filaments  and  by  a  completely  different  color 
pattern;  it  differs  from  C.  kotthtiusi  in  its  shorter 
main  tip  of  preopercular  spine,  shorter  first  spine 
of  first  dorsal  fin,  shorter  caudal  fin  (especially 


FRICKE:   KAIANUS-GROUP  OF  GENUS  CALLIONYMUS 


363 


10mm 


2mm 


2mm 


D 


10mm 


FIGURE  10.  Ccillionymus  kotrhauxi,  ZIM  5535,  holotype,  114.8  mm  SL,  off  Cochin  (India):  (A)  lateral  view;  (B)  left  pre- 
opercular  spine;  (C)  right  preopercular  spine.  ZIM  5536,  paratype,  specimen  I,  male,  81.2  mm  SL,  Cochin:  (D)  first  and 
second  dorsal  fins. 


in  males),  and  various  color  markings  (e.g.,  col- 
or patterns  of  first  and  second  dorsal  fins,  and 
anal  fin,  head,  and  sides  of  body). 

Callionymus  kotthausi  new  name 

(Figure  10) 

Callionymus  ktrianus:  (non  Giinther,  1880)  A i  COCK   1899:74 

(Malabar  Coast,  India,  102  fms  [187  m]). 
Diplof*rammus   imiicus    KOTTHAUS,    1977:40-41,  figs.   423/>, 

424/7,  425  (wsw  of  Cochin,  India). 

MATERIAL  EXAMINED. — Holotype:  ZIM  5535,  9,  114.8  mm 
SL,  A.  Kotthaus,  R/V  METEOR,  10  Feb.  1965,  about  40  km 
wsw  of  Cochin,  India  (09°40'N,  75°38.8'E  to  09°45.3'N, 


75°38.5'E),  211-138  m.  Paratypes:  ZIM  5536,  2  d,  9  9,  71.3- 
87.0  mm  SL;  same  data  as  holotype. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.8-4.1  in  SL);  preopercular  spine  with  a  small 
antrorse  and  one  or  two  large  curved  points,  and 
a  long  main  tip;  second  dorsal  fin  with  a  nearly 
straight  distal  margin;  caudal  fin  without  fila- 
ments; anal  fin  with  a  broad  dark  margin;  sides 
of  body  with  a  row  of  small  distinct  black  blotch- 
es; and  colorless  lower  part  of  caudal  fin. 

DESCRIPTION. — Counts  and   measurements 


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(see  also  Table  1):  D,  IV;  D2  viii.l;  A  viii.l;  P, 
i-ii,17,ii;  P2  1,5;  C  ii,3,i,3,iii. 

Body  elongate,  depressed.  Head  depressed, 
3.8-4.1  in  SL.  Eye  large,  about  2.4  in  head.  Pu- 
pil about  2.8  in  eye.  Branchial  opening  of  normal 
size,  about  2-3  times  in  pupil.  Occipital  region 
with  a  low  bony  protuberance.  Preopercular 
spine  about  1 .4  in  eye  diameter,  with  a  long  up- 
curved  main  tip,  a  small  antrorse  and  one  or  two 
large  curved  points  on  its  dorsal  side,  and  a  large 

antrorse  spine  at  its  base  (formula  1 1;  see 

Fig.  IQfl.C).  Lateral  line  reaching  from  area  be- 
hind eye  to  end  of  median  unbranched  caudal 
fin  ray,  with  a  long  branch  at  its  ventral  side  in 
postorbital  region;  the  line  of  the  opposite  side 
is  interconnected  by  a  transverse  branch  across 
the  occipital  region  and  another  across  the  cau- 
dal peduncle.  Caudal  peduncle  length  about  6.5 
in  SL;  minimal  caudal  peduncle  depth  about  24.0 
in  SL. 

First  spine  of  first  dorsal  fin  long  (longer  than 
first  ray  of  second  dorsal  fin),  but  not  filamen- 
tous. Distal  margin  of  second  dorsal  fin  nearly 
straight.  Anal  fin  beginning  on  the  vertical 
through  third  ray  of  second  dorsal  fin.  Distal 
margin  of  caudal  fin  convex  or  slightly  pointed, 
without  filaments.  Distal  margin  of  pelvic  fin 
convex;  longest  pelvic  fin  ray  reaching  only  to 
anal  papilla  when  laid  back.  Pectoral  fin  reaching 
to  middle  of  third  membrane  of  second  dorsal 
fin  when  laid  back. 

Color  in  alcohol.  Head  and  dorsal  side  of  body 
yellowish  brown;  back  with  some  dark-edged 
whitish  blotches.  Sides  of  body  with  a  row  of 
dark  spots.  Thorax,  belly,  and  lower  parts  of 
body  yellowish  white.  Eye  dark  gray.  Opercu- 
lum  with  dark  spots.  First  dorsal  fin  brownish; 
males  with  one  or  two  black  blotches  surround- 
ed by  white  on  second  and  third  membranes; 
females  with  an  elongate  ocellated  basal  black 
blotch  on  first  to  third  membranes.  Second  dor- 
sal fin  transparent,  distally  darkish,  with  two 
rows  of  elongate  dark  spots.  Distal  margin  of 
anal  fin  black,  tips  of  rays  white.  Caudal  fin 
mostly  colorless,  with  3-4  vertical  rows  of  light 
brown  spots  in  its  upper  part.  Distal  half  of 
fourth  and  fifth  pelvic  fin  rays  dark;  pelvic  fin 
basally  with  irregular  darkish  spots. 

DISTRIBUTION. — Southwest  coast  of  India 
(see  Fig.  2);  138-211  m. 

DISCUSSION. — Kotthaus  (1977)  assigned  Dip- 
logrammus  indicus  to  the  genus  Diplogrammus 


using  Smith's  (1963:549)  key:  "A  skinny  keel 
along  lower  flank  from  tip  of  pelvic  to  caudal 
base,"  in  combination  with  "an  antrorse  spine 
at  base  of  preopercular  spine.1'  The  latter  feature 
is  also  valid  for  species  of  the  genus  Calliony- 
mus,  but  the  specimens  of  "Diplogrammus  in- 
dicus" have  neither  a  skinny  keel  along  lower 
flank  of  body,  as  in  species  of  Diplogrammus, 
nor  any  other  diagnostic  feature  of  Diplogram- 
mus (e.g.,  a  free  flap  of  skin  at  the  operculum, 
a  lateral  line  with  many  branches).  Kotthaus' s 
species  belongs  in  the  genus  Callionymus,  and 
it  posseses  all  features  of  the  kaianus-group  of 
the  subgenus  Callionymus.  The  binomen  Cal- 
lionymus indicus  (Kotthaus,  1977),  however, 
becomes  a  secondary  homonym  of  Callionymus 
indicus  Linnaeus,  1758  (a  Platycephalidae  now 
well  known  as  Platycephalus  indicus)  and  must 
be  replaced. 

ETYMOLOGY. — The  species  is  named  after  Dr. 
A.  Kotthaus,  who  first  described  the  species.  In 
accordance  with  Dr.  Kotthaus,  who  is  presently 
unable  to  create  a  new  name  for  the  species  be- 
cause of  his  health,  I  propose  the  new  name 
Callionymus  kotthausi  to  replace  Callionymus 
indicus  (Kotthaus,  1977). 

Callionymus  africanus  (Kotthaus,  1977) 

(Figure  11) 

Callionymus  kaianus:  (non  Giinther  1880)  NORMAN  1939:73 
(Zanzibar  area);  SMITH  1963  (part):553  (after  Norman). 

Diplogrammus  africanus  KOTTHAUS,  1977(part):38— 40,  figs. 
421,  422,  423«,  424«  (NE  of  Mombasa). 

MATERIAL  EXAMINED.— Holotype:  ZIM  5533,  2,  102.3 
mm  SL,  A.  Kotthaus,  R/V  METEOR,  14  Jan.  1965,  about  180 
naut.  miles  NE  of  Mombasa  (01°18'S,  41°56'E  to  OP19.8'S, 
41°53'E).  Paratypes:  ZIM  5534,  11  6,  19  9,  same  data  as 
holotype. 

Other  specimens:  BMNH  1939.5.24.1422,  1  6,  75.5  mm 
SL,  John  Murray  Exped.,  12  Jan.  1934,  near  Zanzibar 
(5°38'54"S,  39°15'42"E  to  5°40'I8"S,  39°17'36"E);  green  mud 
bottom;  bottom  temperature  15.52°C;  bottom  salinity  35.21%r. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(about  4.0  in  SL);  preopercular  spine  with  a 
small  antrorse  and  two  larger  curved  points  at 
dorsal  side,  and  a  relatively  short  main  tip;  near- 
ly straight  distal  margin  of  second  dorsal  fin;  two 
short  median  caudal  fin  filaments;  first  dorsal  fin 
without  a  filament  and  with  a  normal  (not  in- 
cised) second  membrane;  anal  fin  with  a  narrow 
black  distal  margin;  no  vertically  elongated  dark 
blotches  in  median  part  of  caudal  fin;  back  with 


FRICKE:   KAIAN US-GROUP  OF  GENUS  CALL1ONYMUS 


365 


A 


B 


2mm 


^^•^^•w 

10mm 


FIGURE  11.     Callionymus  africanus,  BMNH  1939.5.24.1422,  male,  75.5  mm  SL,  Zanzibar:  (A)  lateral  view;  (fi)  left  pre- 
opercular  spine. 


dark-bordered  light  spots;  and  pectoral  fin  base 
with  two  transverse  dark  streaks. 

DESCRIPTION. — Counts  and  proportions  (see 
also  Table  1):  D,  IV;  D2  viii.l;  A  viii.l;  P,  i- 
ii,16-20,i-ii;  P2  1,5;  C  ii,3,ii,2,iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.9-4.1  in  SL.  Eye  large,  2.0-2.2  in 
head.  Pupil  relatively  small,  about  3.7  in  eye 
diameter.  Branchial  opening  of  normal  size, 
about  3  in  pupil.  Occipital  region  with  two  low 
bony  protuberances.  Preopercular  spine  1.8  in 
eye  diameter,  with  a  short,  slightly  upcurved 
main  tip,  a  small  antrorse  and  two  large  curved 
points  on  its  dorsal  side,  and  a  large  antrorse 

spine  at  its  base  (formula:   1 1;  see  Fig. 

1  IB).  Lateral  line  reaching  from  area  behind  eye 
to  end  of  third  branched  caudal  fin  ray  (counted 
from  above);  the  line  of  the  opposite  side  is  in- 
terconnected by  a  transverse  branch  across  oc- 
cipital region  and  another  across  dorsal  side  of 
caudal  peduncle.  Lateral  line  with  a  long  branch 
at  its  ventral  side  behind  eye.  Caudal  peduncle 
length  6.0-6.1  in  SL,  minimal  caudal  peduncle 
depth  19.9-21. 3  in  SL. 

First  spine  of  first  dorsal  fin  somewhat  longer 
than  first  ray  of  second  dorsal  fin,  but  not  fila- 
mentous. Distal  margin  of  second  dorsal  fin 
nearly  straight.  Anal  fin  beginning  on  the  vertical 
through  second  ray  of  second  dorsal  fin.  Distal 


margin  of  caudal  fin  convex;  males  with  two  rel- 
atively short  median  filaments,  females  without 
filaments.  Distal  margin  of  pelvic  fin  convex; 
longest  pelvic  fin  ray  reaching  to  base  of  first 
anal  fin  ray  when  laid  back.  Pectoral  fin  reaching 
to  fourth  ray  of  second  dorsal  fin  when  laid  back. 

Color  in  alcohol.  Head  and  body  brown;  ven- 
tral side  of  body  lighter,  belly  and  thorax  white. 
A  row  of  paired  dark  brown  blotches  on  side  of 
body.  Back  with  light  spots  bordered  with  dark 
brown.  Pectoral  fin  base  with  two  transverse 
dark  lines.  Eye  dark  blue,  dorsally  with  dark 
brown  blotches.  First  dorsal  fin  brown;  a  large 
white-edged  black  blotch  basally  on  third  mem- 
brane, a  basal  and  a  distal  branch  of  which 
reaches  to  second  spine;  occasionally,  also  with 
a  small  distal  black  blotch  on  third  membrane. 
Second  dorsal  fin  with  three  rows  of  white  and 
two  rows  of  darkish  spots;  these  rows  alternate- 
ly arranged.  Anal  fin  pale,  with  a  narrow  black 
distal  margin.  Lower  part  of  caudal  fin  with  a 
broad  curved  dark  bar,  upper  part  scattered  with 
dusky  spots.  Distal  parts  of  fourth  and  fifth  pel- 
vic fin  rays  dark;  upper  part  of  pectoral  fin  with 
three  vertical  rows  of  dark  spots. 

SEXUAL  DIMORPHISM. — As  described  by 
Kotthaus  (1977:40). 

DISTRIBUTION. — Known  from  Zanzibar  and 
southern  Somalia  (see  Fig.  1),  possibly  distrib- 


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uted  along  entire  east  African  coast;  212  m,  mud 
bottom. 

DISCUSSION. — The  differences  from  Callio- 
nymus  regani  are  discussed  in  the  description  of 
that  species.  The  two  paratypes  described  by 
Kotthaus  (1977:39)  from  the  southern  Red  Sea 
are  small  specimens  of  Callionymus  bentuviai. 

Callionymus  bentuviai  new  species 

(Figure  12) 

Diplogrammus  africanus   KOTTHAUS,   1977  (part):39  (s  Red 
Sea). 

MATERIAL  EXAMINED. — Holotype:  HUJF  9935,  87.8  mm 
SL,  A.  Ben-Tuvia,  1957,  Eritrea  (Ethiopia;  s  Red  Sea).  Para- 
types:  HUJF  8068,  2  spec.,  79.7-85.2  mm  SL,  same  data  as 
holotype;  ZIM  5532,  2  spec.,  34.2-46.9  mm  SL,  A.  Kotthaus, 
R/V  METEOR,  6  Dec.  1964,  s  Red  Sea,  70-75  m. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.6-4.0  in  SL);  preopercular  spine  with  a  small 
antrorse  and  two  larger  curved  points  at  its  dor- 
sal side,  and  a  relatively  long  main  tip;  second 
dorsal  fin  with  a  nearly  straight  distal  margin; 
two  very  long  median  caudal  fin  filaments;  first 
dorsal  fin  without  a  filament  and  with  a  deeply 
incised  second  membrane;  and  second  dorsal  fin 
with  vertical  dark  streaks. 

DESCRIPTION. — Counts  and  proportions  (see 
also  Table  1):  D,  IV;  D2  viii,l;  A  viii.l;  P,  i- 
ii,15-17,i-ii;  P2  1,5;  C  ii,2-4,i-ii,2-3,ii-iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.6-4.0  in  SL.  Eye  large,  2.15-2.85  in 
head.  Pupil  relatively  small,  about  2.7  in  eye. 
Branchial  opening  of  normal  size,  about  1.5  in 
pupil.  Occipital  region  with  two  low  bony  pro- 
tuberances. Preopercular  spine  1.2-1.6  in  eye 
diameter,  with  a  long  slightly  upcurved  main  tip, 
a  small  antrorse  and  two  larger  curved  points  on 
its  dorsal  side,  and  a  large  antrorse  spine  at  its 
base  (formula:  1 1;  see  Fig.  12C,£>).  Lat- 
eral line  reaching  from  area  behind  eye  to  end 
of  third  branched  caudal  fin  ray  (counted  from 
above);  the  line  of  the  opposite  side  is  intercon- 
nected by  a  transverse  branch  across  occipital 
region  and  another  across  caudal  peduncle.  Cau- 
dal peduncle  length  4.9-5.8  in  SL,  minimal  cau- 
dal peduncle  depth  23.1-24.9  in  SL. 

First  spine  of  first  dorsal  fin  longer  than  first 
ray  of  second  dorsal  fin,  but  not  filamentous. 
Second  membrane  of  first  dorsal  fin  deeply  in- 
cised (not  in  young  specimens).  Distal  margin  of 
second  dorsal  fin  nearly  straight.  Anal  fin  begin- 
ning on  the  vertical  through  midbase  of  second 


membrane  of  second  dorsal  fin.  Distal  margin  of 
caudal  fin  convex,  with  two  very  long  median 
filaments.  Distal  margin  of  pelvic  fin  mostly  con- 
vex; longest  ray  reaching  to  middle  of  second 
membrane  of  anal  fin  when  laid  back.  Pectoral 
fin  reaching  to  fourth  ray  of  second  dorsal  fin 
when  laid  back. 

Color  in  alcohol.  Head  and  body  dark  brown; 
ventral  side  of  body  dark  brown,  belly  whitish. 
Back  nearly  monochromatic,  with  very  few 
small  dark  spots.  A  row  of  small  pale  blotches 
along  side  of  body,  occasionally  also  a  row  of 
two  groups  of  three  dark  spots  each.  Many  black 
spots  on  operculum  and  at  pectoral  fin  base. 
First  dorsal  fin  light  brown,  with  two 
darkish  transverse  lines  distally  between  first 
and  third  spines.  A  large,  elongate,  curved  black 
blotch  basally  from  first  to  fourth  spine,  mainly 
on  third  membrane.  Another  black  blotch  dis- 
tally on  third  membrane.  Second  dorsal  fin  pale, 
with  nine  vertical  dark  streaks.  Anal  fin  dark 
brown,  with  a  black  distal  margin.  Caudal  fin 
pale;  dorsal  part  with  a  narrow  curved  dark  line, 
ventral  part  with  a  broad  curved  dark  bar.  Pelvic 
fin  pale;  upper  one-third  with  three  vertical  rows 
of  dark  spots. 

DISTRIBUTION. — Known  only  from  the  south- 
ern Red  Sea  (see  Fig.  2);  70-75  m. 

DISCUSSION. — Callionymus  bentuviai  differs 
from  all  other  species  of  the  kaianus -group  in 
its  deeply  incised  second  membrane  of  the  first 
dorsal  fin,  the  absence  of  a  dorsal  fin  filament  in 
combination  with  two  very  long  median  caudal 
fin  filaments,  and  an  unusual  color  pattern  (e.g., 
in  the  dorsal  fins). 

ETYMOLOGY. — This  species  is  named  for  Pro- 
fessor Adam  Ben-Tuvia  (Hebrew  University  of 
Jerusalem),  who  collected  the  holotype  and  sent 
it  to  me  for  examination. 

Callionymus  ochiaii  new  species 

(Figure  13) 

Callionymux  kaianus:  (non  Giinther,  1880)  OCHIAI,  ARAGA, 
AND  NAKAJIMA  1955: 1 1 1-113,  figs.  8-10,  table  6  (various  s 
Japan  localities);  MATSUBARA  1955:713  (after  Ochiai,  Ara- 
ga,  and  Nakajima);  KAMOHARA  1964:90  (Kochi  Pref.  to 
Kagoshima:  deep-sea  bottom,  very  rare);  MASUDA,  ARAGA, 
AND  YOSHINO  1975:261,  pi.  847)  (Kumano  Bay  south). 

MATERIAL  EXAMINED.— Holotype:  FAKU  23261,  $,  95.0 
mm  SL,  1-3  Sep.  1954,  Shibushi,  Kagoshima  Pref.,  Japan. 
Paratypes:  FAKU  23257-23260  and  FAKU  23275,  2  6,  3  9, 
77.8-116.8  mm  SL,  same  data  as  holotype. 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 


FRICKE:   KAIAN US-GROUP  OF  GENUS  CALLIONYMUS 


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10  mm 


C  2mm  D 

FIGURE  12.     Ctillionymus  bentuviai,  HUJF  9935,  holotype,  87.8  mm  SL,  Eritrea:  (A)  lateral  view;  (B)  dorsal  view;  (C)  left 
preopercular  spine:  (D)  right  preopercular  spine. 


(3.5-4.4  in  SL);  preopercular  spine  with  a  small 
antrorse  and  two  larger  curved  points  at  its  dor- 
sal side,  and  a  relatively  long  main  tip;  nearly 
straight  distal  margin  of  second  dorsal  fin;  one 
or  two  short  median  caudal  fin  filaments;  first 
dorsal  fin  with  a  long  filament  (males)  or  a  fila- 
ment of  median  size  (females);  anal  fin  without 
a  distal  dark  margin;  and  long,  slender  main  tip 
of  preopercular  spine,  which  is  longer  than  the 
longest  point  on  its  dorsal  edge. 


DESCRIPTION. — Counts  and  proportions  (see 
also  Table  2):  D,  IV;  D2  viii.l;  A  viii.l;  P,  ii,16- 
18,i;  P2  1,5;  C  ii,3(-4),i,(2-)3,iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.5-4.4  in  SL.  Eye  large,  2.3-3.0  in 
head.  Pupil  relatively  small,  about  3.65  in  eye 
diameter.  Branchial  opening  of  normal  size, 
about  2.5  in  pupil.  Occipital  region  with  two  low 
bony  ridges.  Preopercular  spine  1.3-1.8  in  eye 
diameter,  with  a  long,  slightly  upcurved  main 


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FIGURE  13.     Callionymux  ochiuii,  holotype,  FAKU  23261,  male,  95.0  mm  SL,  Shibushi  (Japan):  (A)  lateral  view;  (B)  left 
preopercular  spine.  Paratype,  FAKU  23258,  female,  116.8  mm  SL,  Shibushi  (Japan):  (C)  first  dorsal  fin. 


tip,  a  small  antrorse  and  two  larger  curved 
points  at  its  dorsal  side,  and  a  large  antrorse 

spine  at  its  base  (formula:   1 1;  see  Fig. 

135).  Lateral  line  reaching  from  area  behind  eye 
(from  where  a  branch  runs  down  to  base  of  pre- 
opercular spine)  to  end  of  third  branched  caudal 
fin  ray  (counted  from  above);  the  line  of  the  op- 
posite side  is  interconnected  by  a  transverse 
branch  across  occipital  region  and  another 
across  dorsal  part  of  caudal  peduncle.  Caudal 
peduncle  length  5.3-6.1  in  SL;  minimal  caudal 
peduncle  depth  24.3-27.8  in  SL. 

First  spine  of  first  dorsal  fin  filamentous  (fil- 
ament in  females  relatively  short).  Second  mem- 
brane of  first  dorsal  fin  not  incised.  Distal  margin 
of  second  dorsal  fin  nearly  straight.  Anal  fin  be- 
ginning on  a  vertical  through  midbase  of  second 


membrane  of  second  dorsal  fin.  Distal  margin  of 
caudal  fin  irregular,  with  two  short  median  fila- 
ments. Distal  margin  of  pelvic  fin  convex;  long- 
est pelvic  fin  ray  reaching  to  midbase  of  first 
anal  fin  membrane.  Pectoral  fin  reaching  to 
fourth  ray  of  second  dorsal  fin  when  laid  back. 
Color  in  alcohol.  Body  brownish  yellow 
above,  whitish  below.  Back  with  minute  black- 
ish spots,  forming  rings  and  blotches.  A  row  of 
dark  brown  blotches  along  sides  of  body.  Pec- 
toral fin  base  with  a  dark  area.  Head  with  some 
irregular  dark  spots  and  blotches.  First  dorsal 
fin  gray,  with  a  black  blotch  basally  or  centrally 
on  third  membrane,  a  basal  branch  of  which  oc- 
casionally reaches  to  second  spine.  Second  dor- 
sal fin  gray,  with  large  irregular  dark  blotches 
and  submarginal  dark  bands.  Caudal  fin  pale  ex- 


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369 


cept  for  the  darker  lower  part  where  the  fin  is 
scattered  with  several  pale  pearl-white  round 
spots.  Anal  fin  dusky,  sometimes  indefinitely  but 
broadly  edged  with  brown  (but  not  with  black). 
Pectoral  fin  uniformly  pale;  pelvic  fin  pale,  outer 
posterior  margin  more  or  less  dark. 

DISTRIBUTION. — Southern  Japan  (see  Fig.  1); 
about  100  m,  sand  and  mud  bottoms. 

DISCUSSION. — From  the  most  similar  species, 
C.  moretonensis,  C.  ochiaii  differs  in  having  a 
somewhat  shorter  main  tip  of  preopercular 
spine,  an  irregular  distal  margin  of  caudal  fin, 
and  several  different  color  markings  (especially 
the  absence  of  a  distal  black  band  in  the  anal  fin; 
the  color  patterns  of  the  caudal  and  second  dor- 
sal fins  and  of  the  pectoral  fin  base;  the  body 
color  pattern,  etc.).  Callionymus  ochiaii  differs 
from  C.  kaianus  in  the  shape  of  the  largest  point 
on  the  dorsal  side  of  the  preopercular  spine,  in 
the  length  of  the  filament  in  the  first  dorsal  fin, 
and  in  various  color  markings. 

The  first  dorsal  fin  filament  of  the  male  spec- 
imen figured  in  Ochiai,  Araga,  and  Nakajima 
(1955:fig.  8)  is  very  short.  Because  the  specimen 
(91.0  mm  SL)  is  nearly  as  long  as  the  holotype 
(95.0  mm  SL),  the  filament  might  have  been  bro- 
ken in  that  specimen. 

ETYMOLOGY. — This  new  species  is  named  for 
Dr.  Akira  Ochiai,  who  first  described  the  species 
under  the  name  Callionymus  kaianus. 

Callionymus  formosanus  new  species 

(Figure  14) 

"!Callion\mus  kaianus:  (non  Giinther,  1880)  CHU  1957:22  (Pes- 
cadores Islands). 

MATERIAL  EXAMINED.— Holotype:  CAS  46972,  <S ,  104.0 
mm  SL,  F.  B.  Steiner,  Apr.  1971,  Formosa  Str.  (25°N,  120°E), 
approximately  90  m. 

DIAGNOSIS. — A  Callionymus  of  the  kaitinus- 
group  with  a  small  branchial  opening;  short  head 
(about  4.5  in  SL);  preopercular  spine  with  a 
small  antrorse  and  two  larger  curved  points  at 
dorsal  side;  relatively  high  second  dorsal  and 
anal  fins  with  convex  distal  margins;  filamentous 
first  spine  of  first  dorsal  fin;  large  black  blotch 
nearly  on  entire  third  membrane  of  first  dorsal 
fin;  anal  fin  with  a  distal  black  streak  on  each 
membrane;  distal  two-thirds  of  anal  fin  brown; 
and  white  anal  fin  ray  tips. 

DESCRIPTION. — Counts  and  proportions  of  the 
holotype  (see  also  Table  2):  D,  IV;  D2  viii,l;  A 
viii.l;  P,  ii,16-17,ii;  P2  1,5;  C  ii,3,ii,2,iii. 

Body  elongate  and  depressed.   Head  de- 


pressed, about  4.5  in  SL.  Eye  large,  about  2.3 
in  head.  Pupil  relatively  small,  about  3.4  in  eye 
diameter.  Branchial  opening  of  normal  size, 
about  2.5  in  pupil.  Occipital  region  with  two  low 
bony  ridges.  Preopercular  spine  about  1.7  in  eye 
diameter,  with  a  slightly  upcurved  main  tip  of 
medium  size,  a  small  antrorse  and  two  larger 
curved  points  at  its  dorsal  side,  and  a  large  an- 
trorse spine  at  its  base  (formula:  1 1;  see 

Fig.  14B).  Lateral  line  reaching  from  area  behind 
eye  (from  where  a  long  branch  runs  downwards) 
to  middle  of  upper  median  unbranched  caudal 
fin  ray;  the  line  of  the  opposite  side  is  intercon- 
nected by  a  transverse  branch  across  occipital 
region  and  another  across  dorsal  side  of  caudal 
peduncle.  Caudal  peduncle  length  4.9  in  SL, 
minimal  caudal  peduncle  depth  13.0  in  SL. 

First  spine  of  first  dorsal  fin  filamentous.  Sec- 
ond dorsal  fin  relatively  high,  distal  margin  con- 
vex. Anal  fin  beginning  on  a  vertical  through 
midbase  of  first  membrane  of  second  dorsal  fin. 
Distal  margin  of  caudal  fin  convex,  with  one  rel- 
atively long  filament.  Distal  margin  of  pelvic  fin 
convex;  longest  pelvic  fin  ray  reaching  to  mid- 
base  of  first  membrane  of  anal  fin.  Pectoral  fin 
reaching  to  third  ray  of  second  dorsal  fin  when 
laid  back. 

Color  in  alcohol.  Body  dark  brown,  belly 
whitish.  Side  of  body  with  a  row  of  irregular 
blackish  blotches.  Head  dark  brown,  with  some 
lighter  spots.  A  vertical  dark  streak  under  the 
eye.  A  dark  blotch  at  pectoral  fin  base.  Back 
and  sides  of  body  with  whitish  dark-edged 
blotches.  First  dorsal  fin  pale,  with  a  large  dark 
blotch  on  third  membrane  and  three  white 
blotches  on  first  and  second  membranes.  Second 
membrane  with  a  distal  black  margin.  Second 
dorsal  fin  pale,  with  a  vertical  white  streak  on 
each  fin  ray.  Distal  margin  of  first  three  rays  of 
second  dorsal  fin  black.  Anal  fin  pale,  distal  two- 
thirds  brownish,  with  a  distal  black  streak  on 
each  membrane;  tips  of  fin  rays  white.  Caudal 
fin  pale,  with  a  vertical  row  of  dark  spots;  lower 
margin  and  lower  distal  margin  dark.  Pelvic  fin 
with  irregular  darkish  spots  and  a  dark  distal 
margin  of  third,  fourth,  and  fifth  rays.  Upper 
part  of  pectoral  fin  with  three  vertical  rows  of 
dark  spots. 

DISTRIBUTION. — Known  only  from  Formosa 
Strait  (see  Fig.  2):  about  90  m. 

DISCUSSION. — This  interesting  new  species 
belongs  to  the  subgroup  of  species  with  a  high 
second  dorsal  fin  in  the  kaianus  -group.  It  seems 


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FIGURE  14.     Callionymusformosanus,  holotype,  CAS  46972,  male,  104.0  mm  SL,  Formosa  Strait:  (A)  lateral  view;  (B)  left 
preopercular  spine. 


to  be  a  primitive  member  of  that  subgroup  be- 
cause the  second  dorsal  fin  is  still  relatively  low. 

The  specimen  recorded  as  Callionymus 
kaianus  by  Chu  (1957)  from  Pescadores  Islands 
(southern  part  of  Formosa  Strait)  is  probably 
this  species. 

ETYMOLOGY. — This  new  species  is  named  af- 
ter its  type-locality  (Formosa  Strait). 

Callionymus  guentheri  new  species 

(Figures  15-17) 

Callionymus  curvicornis:  (non  Valenciennes,  1837)  GUNTHER 
1880:  53  (Philippines,  82  fms  [150  m]);  HERRE  1953:777  (af- 
ter Giinther). 

MATERIAL  EXAMINED.— Holotype:  BMNH  1879.5.14.567, 
1,  87.3  mm  SL,  CHALLENGER  Exped.,  26  Oct.  1874,  w  of 
Zamboanga,  Philippine  Is.  (7°03'N,  121°48'E;  entrance  from 
Sulu  Sea  into  Basilan  Str.,  about  7  miles  w  of  Mindanao  I., 
and  to  the  NW  of  Caldera  Pt.),  82  fms  (150  m).  Paratypes  (all 
from  the  Philippines  collected  by  J.  E.  Norton,  1966):  CAS 
46966,  4  spec.,  3  Nov.,  Sandoval  Pt.,  Catanauan,  Quezon,  70- 
78  fms  (128-143  m);  CAS  32897,  11  spec.  (7  6 ,  4  9),  51.2- 
128.0  mm  SL,  27  June,  Lemery  Town,  Balayan  Bay,  Batan- 
gas,  Luzon,  90-85  fms  (165-155  m);  CAS  33879,  1  9,  112.8 
mm  SL,  15  Dec.,  N  of  San  Andres  I.,  Marinduque,  151-158 
fms  (276-289  m);  CAS  34286,  3  6,  100.0-108.3  mm  SL,  14 


Dec.,  NW  of  San  Andres  I.,  Marinduque,  137-139  fms  (250- 
254  m);  CAS  34197,  1  d,  8  9,  90.2-129.3  mm  SL,  24  Nov., 
Siburio  Pt.,  Ragay  Gulf,  Camarines  Sur  Prov.,  319-324  fms 
(583-593  m);  CAS  32905,  4  d,  3  9,  69.3-117.2  mm  SL,  15 
Nov.,  Buri  Pt.,  Ragay  Gulf,  Camarines  Sur  Prov.,  304-309 
fms  (556-566  m);  CAS  34426,  1  9,  84.1  mm  SL,  4  July,  s  of 
Bauan  Town,  Batangas  Bay,  Batangas  Prov.,  90-88  fms  (165- 
161  m);  CAS  34278,  4  9,  89.9-124.5  mm  SL,  11  Dec.,  N  of 
Melchor  I.,  Marinduque,  120-126  fms  (219-231  m):  CAS 
33864,  3  9,  90.0-127.3  mm  SL,  23  Nov.,  Caurusan  Pt.,  Ragay 
Gulf,  Camarines  Sur  Prov.,  302-308  fms  (552-564  m);  CAS 
32916,  5  9,  91.0-118.2  mm  SL,  11  Nov.,  Pusgo  Pt.,  Ragay 
Gulf,  Quezon,  Luzon  I.,  60-67  fms  (110-123  m);  CAS  33703, 
1  9,  122.0  mm  SL,  25  June,  s  of  Barrio  Nomong  Casto,  Ba- 
layan Bay,  105-100  fms  (183-192  m);  CAS  34401,  1  9,  117.7 
mm  SL,  24  June,  SE  of  Calaca  Town,  Balayan  Bay,  65-55  fms 
(119-100  m);  CAS  32997,  1  9,  107.5mm  SL,  15  June,  Batan- 
gas, Balayan  Bay,  s  of  Barrio  Sinisian,  95-99  fms  (174-181 
m);  CAS  34190,  1  d  (113.2  mm  SL),  1  9  (99.7  mm  SL),  6 
Nov.,  SE  of  Alibijaban  I.,  Ragay  Gulf,  81-88  fms  (148-161  m); 
CAS  34272,  1  d,  108.0  mm  SL,  14  Nov.,  Nagas  Pt.,  Ragay 
Gulf,  297-299  fms  (543-547  m);  CAS  34154,  7  9,  91.1-129.2 
mm  SL,  3  Sep.,  N  of  San  Andres  Pt.,  Marinduque,  108-112 
fms  (197-205  m);  CAS  32668,  2  9 ,  94.5-102.0  mm  SL,  26  July, 
SE  of  Talaga,  Batangas  Bay,  138-131  fms  (253-240  m):  CAS 
32801,  1  d,  91.9  mm  SL,  25  Aug.,  NE  of  Salomague  I.,  Mar- 
induque, 142-150  fms  (260-275  m):  CAS  34205,  1  9,  127.0 
mm  SL,  10  Dec.,  NW  of  Baltazar  I.,  Marinduque,  142-150 fms 
(260-275  m);  CAS  34468,  1  9  ,  102.8  mm  SL,  27  July,  Lemery 


FRICKE:   KAIANUS-GROVP  OF  GENUS  CALLIONYMUS 


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10mm 


FIGURE  15.     Callionymus  guentheri,  holotype,  BMNH  1879.5.14.567,  female,  87.3  mm  SL,  off  Zamboanga,  Philippines:  (A) 
lateral  view;  (B)  left  preopercular  spine. 


Town,  Balayan  Bay,  85-90  fms  (155-164  m);  CAS  34074,  1 
9,  123.2  mm  SL,  N  of  Sayao  Bay,  Marinduque,  61-70  fms 
(111-128  m);  CAS  33067,  1  9,  125.2  mm  SL,  18  July,  s  of 
Barrio  Salong,  Luzon  I.,  120-114  fms  (220-208  m). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.9-4.2  in  SL);  preopercular  spine  with  a  small 
antrorse  and  two  larger  curved  points  on  dorsal 
side;  very  high  second  dorsal  fin  with  a  convex 
distal  margin  (males)  or  a  relatively  low  second 
dorsal  fin  with  a  nearly  straight  distal  margin 
(females);  first  dorsal  fin  without  a  filament  (or 
with  a  very  short  one  in  smaller  specimens);  long 
and  slender  main  tip  of  preopercular  spine;  anal 
fin  with  a  black  distal  half  and  white  fin  ray  tips; 
regular  distal  margin  of  caudal  fin,  with  filaments 
only  in  small  specimens  (caudal  fin  elongate  in 
adults);  and  relatively  large  central  black  blotch 
on  third  membrane  of  first  dorsal  fin. 

DESCRIPTION. — Counts  and  proportions  of  the 
holotype  (female)  (see  also  Table  2):  D,  IV;  D2 
viii.l;  A  viii.l;  P,  i-ii,17-19,i-ii;  P2  1,5;  C  ii,2- 
3,i-ii,2-4,iii. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.9-4.2  in  SL.  Eye  large,  2.3-2.6  in 


head.  Pupil  of  normal  size,  2.7-3.5  in  eye  di- 
ameter. Branchial  opening  of  normal  size,  about 
2.0  in  pupil.  Occipital  region  with  two  low  bony 
ridges.  Preopercular  spine  1.5-1.7  in  eye  diam- 
eter, with  a  slightly  upcurved  main  tip  which  is 
relatively  long  or  of  medium  size,  a  small  an- 
trorse and  two  larger  curved  points  on  its  dorsal 
side,  and  a  large  antrorse  spine  at  its  base  (for- 
mula: 1 — ^ — 1;  see  Fig.  155,  Fig.  16fl).  The 
two  large  spines  on  the  dorsal  side  of  the  pre- 
opercular spine  of  the  holotype  possess  small 
basal  hooks  (see  Fig.  15B).  Lateral  line  usually 
reaching  from  area  behind  eye  to  end  of  third 
branched  caudal  fin  ray  (counted  from  above); 
because  of  the  different  caudal  fin  formula  of  the 
holotype  (C  ii,2,i,4,iii  instead  of  C  ii,3,ii,2,iii), 
the  lateral  line  reaches  to  middle  of  median  un- 
branched  caudal  fin  ray  in  that  specimen.  The 
lateral  line  of  the  opposite  side  is  interconnected 
by  a  transverse  branch  across  occipital  region 
and  another  across  dorsal  side  of  caudal  pedun- 
cle. Caudal  peduncle  length  5.1-6.0  in  SL,  min- 
imal caudal  peduncle  depth  23.0-26.0  in  SL. 

First  spine  of  first  dorsal  fin  somewhat  longer 
than  first  ray  of  second  dorsal  fin  but  not  fila- 


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FIGURE  16.     Callionymus  guentheri,  paratype,  CAS  32897  (specimen  1),  male,  128.0  mm  SL,  Balayan  Bay,  Philippines:  (A) 
lateral  view;  (B)  left  preopercular  spine. 


mentous  (only  occasionally  in  small  specimens 
a  very  short  filament).  Second  dorsal  fin  in  males 
relatively  high,  with  a  convex  distal  margin,  in 
females  lower,  with  a  nearly  straight  distal  mar- 
gin. Anal  fin  beginning  on  the  vertical  through 
base  of  second  ray  of  second  dorsal  fin.  Distal 
margin  of  caudal  fin  in  small  specimens  convex, 
with  one  or  two  long  filaments;  in  larger  speci- 
mens all  median  caudal  fin  rays  elongate  (the 
two  median  rays  longest),  but  not  filamentous; 
even  the  longest  rays  are  connected  by  mem- 
branes. Distal  margin  of  pelvic  fin  convex;  long- 
est pelvic  fin  ray  reaching  to  midbase  of  first  or 
second  membrane  of  anal  fin.  Pectoral  fin  reach- 
ing to  fourth  or  fifth  ray  of  second  dorsal  fin 
when  laid  back. 

Color  in  alcohol.  Head  and  body  brown.  One 
or  two  rows  of  large,  irregular  dark  blotches 
along  side  of  body.  A  large  dark  area  on  pectoral 
fin  base.  Back  with  dark  brown  semicircular  lines 
in  groups  which  are  bordered  by  white  on  one 
side.  Eye  dark  gray.  Belly  and  thorax  white; 
lower  side  of  body  brown.  Head  with  few  irreg- 
ular darkish  spots.  First  dorsal  fin  light  brown, 


with  five  brownish  cross-lines  in  the  anterior 
part;  a  dark  blotch  on  third  membrane  distally 
or  centrally.  Second  dorsal  fin  pale,  females  with 
a  basal  and  a  median  row  of  dark  spots  and  some 
curved  white  lines  (see  Fig.  15A);  males  with 
vertical  darkish  streaks  (see  Fig.  1M).  Distal 
half  of  anal  fin  black;  tips  of  fin  rays  white.  Cau- 
dal fin  in  females  with  a  broad  dark  band  in  the 
lower  and  lower  distal  parts  and  two  median 
short  transverse  dark  bands;  in  the  upper  part 
some  irregular  dark  spots.  Males  have  the  same 
caudal  fin  color  pattern,  but  less  intense.  Pec- 
toral fin  dorsally  with  two  rows  of  dark  spots. 
Distal  three-fourths  of  pelvic  fin  darkish,  with 
irregular  dark  brown  spots  and  blotches. 

DISTRIBUTION. — Known  from  various  locali- 
ties in  the  Philippine  Islands  (southern  coast  of 
Luzon  to  northern  coast  of  Mindanao;  see  Fig. 
2);  100-593  m,  on  mud  bottoms. 

DISCUSSION. — The  differences  between  C. 
guentheri,  C.  whiteheadi,  and  C.  kaianus  have 
been  discussed  in  the  descriptions  of  the  last  two 
species.  Callionymus  guentheri  differs  from  C. 
regani  in  the  caudal  fin  shape,  the  high  second 


FRICKE:   KA1ANUS-GROUP  OF  GENUS  CALLIONYMUS 


373 


c 


10mm 


10mm 


D 


10mm 


10mm 


FIGURE  17.  Callionymux  t>uentheri,  paratypes,  CAS  32897,  Balayan  Bay,  Philippines.  Female  (specimen  2),  108.8  mm  SL: 
(A)  first  and  second  dorsal  fins;  (B)  caudal  fin.  Male  (specimen  4),  77.6  mm  SL:  (C)  first  and  second  dorsal  fins;  (/))  caudal  fin. 
Male  (specimen  10),  85.3  mm  SL:  (E)  first  and  second  dorsal  fins;  (F)  caudal  fin. 


dorsal  fin  (in  males),  and  various  color  markings; 
from  C.  ochiaii  in  the  caudal  fin  shape,  the  lack 
of  a  dorsal  fin  filament  (in  males),  the  high  sec- 
ond dorsal  fin  (in  males),  and  several  color 
markings. 

In  the  extensive  collections  of  J.  E.  Norton 
(paratypes  of  C.  guentheri),  a  great  depth  range 
can  be  found  (100-593  m);  593  m  is  the  greatest 
depth  recorded  for  any  species  of  the  kaianus- 
group.  Because  few  specimens  are  known  of 
most  species  of  that  group,  it  is  possible  that  in 
the  future,  specimens  of  other  species  will  be 
recorded  from  comparable  depths. 

ETYMOLOGY. — This  new  species  is  named  af- 
ter Dr.  Albert  Giinther,  who,  in  1880,  reported 
the  first  specimen  under  the  name  Callionymus 
curvicornis. 


Callionymus  altipinnis  new  species 

(Figure  18) 

Callionymus  ktiianu.i:  (non  Giinther,    1880)  CHU  ET  AL. 

1962:723-724,  fig.   585  (South  China  Sea);   BESEDNOV 

1968:63  (Gulf  of  Tonkin). 
Callionymus  huguenini:  (non  Sleeker,  1858)  SHEN  1964:202- 

203,  fig.  2  (Hong  Kong). 

MATERIAL  EXAMINED.— Holotype:  MSL0001,  6,  123.8mm 
SL,  S.-C.  Shen,  Hong  Kong  Fish  Market.  Paratypes:  CAS 
46967,  10  spec.,  104.1-144.5  mm  SL,  R.  L.  Bolin,  23  July 
1958,  South  China  Sea,  E  of  Hainan  (20°32'N,  1 12°45'30"E); 
CAS  46968,  1  spec.,  135.9  mm  SL,  R.  L.  Bolin,  21  July  1958, 
off  Tungku  Pt.,  Hainan  (19°3rN,  1 11°24'30"E);  CAS  46969, 
1  spec.,  124.8  mm  SL,  R.  L.  Bolin,  23  July  1958,  South  China 
Sea,  E  of  Hainan  (20°32'N,  112°51'E). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(about  4.2  in  SL);  preopercular  spine  with  a  rel- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


A 


10mm 


B 


2mm 


FIGURE  18.     Callionymux  altipinnis,  holotype,  MSL  0001,  male,  123.8  mm  SL,  Hong  Kong:  (A)  lateral  view;  (B)  left  pre- 
opercular  spine. 


atively  short  main  tip;  small  antrorse  and  one 
(rarely  two)  larger  curved  points  at  dorsal  side; 
very  high  second  dorsal  and  anal  fins  with  con- 
vex distal  margins;  filamentous  first  spine  of  first 
dorsal  fin;  small  distal  black  blotch  on  third 
membrane  of  first  dorsal  fin;  anal  fin  with  a  black 
distal  margin  (including  tips  of  fin  rays),  but 
without  a  distal  black  streak  on  each  membrane 
surrounded  by  dark  brown;  and  one  or  two  cau- 
dal fin  filaments  of  medium  size. 

DESCRIPTION. — Counts  and  proportions  of  the 
holotype  (see  also  Table  2):  D,  IV;  D2  viii.l;  A 
viii.l;  P,  i,14-15,ii;  P2  1,5;  C  i,3,ii,3,ii. 

Body  elongate  and  depressed.  Head  de- 
pressed, about  4.2  in  SL.  Eye  large,  about  2.5 
in  head.  Pupil  of  normal  size,  about  3.8  in  eye. 
Branchial  opening  about  3  in  pupil.  Occipital  re- 
gion with  two  low  bony  ridges.  Preopercular 
spine  about  1.7  in  eye  diameter,  with  a  relatively 
short,  slightly  upcurved  main  tip,  a  small  an- 
trorse and  one  (rarely  two)  larger  points  on  its 
dorsal  side,  and  a  large  antrorse  spine  at  its  base 


(formula:  1 


2(-3) 


1;  see  Fig.  185).  Lateral  line 


reaching  from  area  behind  eye  to  middle  of  up- 
per median  unbranched  caudal  fin  ray;  the  line 
of  the  opposite  side  is  interconnected  by  a  trans- 
verse branch  across  occipital  region  and  another 
across  dorsal  side  of  caudal  peduncle.  Caudal 
peduncle  length  about  5.0  in  SL;  minimal  caudal 
peduncle  depth  about  25.0  in  SL. 

First  spine  of  first  dorsal  fin  filamentous.  Sec- 
ond dorsal  fin  very  high,  distal  margin  convex. 
Anal  fin  begins  on  vertical  through  middle  of 
second  membrane  of  second  dorsal  fin.  Distal 
margin  of  pelvic  fin  convex;  longest  pelvic  fin 
ray  reaching  nearly  to  base  of  first  anal  fin  ray. 
Pectoral  fin  reaching  to  middle  of  third  mem- 
brane of  second  dorsal  fin  when  laid  back.  Distal 
margin  of  caudal  fin  somewhat  pointed,  with  one 
or  two  filaments. 

Color  in  alcohol.  Head,  body,  and  fins  pale 
except  for  a  distal  black  blotch  on  third  mem- 
brane of  first  dorsal  fin,  a  row  of  dark  blotches 
along  side  of  body,  eyes  grayish,  a  black  distal 


FRICKE:   KAIAN US  -GROUP  OF  GENUS  CALLIONYMUS 


375 


margin  of  anal  fin,  and  a  darkish  area  on  pectoral 
fin  base. 

DISTRIBUTION. — Known  only  from  the  South 
China  Sea  off  China,  from  Hong  Kong  (type- 
locality)  to  the  Gulf  of  Tonkin;  island  of  Hainan 
(see  Fig.  2).  The  exact  collecting  depth  of  spec- 
imens of  C.  altipinnis  is  not  known,  but  the 
species  should  occur  at  about  the  same  depth  as 
other  species  of  the  kaianus -group. 

DISCUSSION. — Callionymus  altipinnis  differs 
from  C.  sokonumeri  in  the  filamentous  first 
spine  of  the  first  dorsal  fin,  the  shorter  caudal 
fin  filaments,  and  various  color  markings. 

The  record  of  C.  huguenini  Bleeker  (by  Shen 
1964)  from  Hong  Kong  is  based  on  a  misidenti- 
fied  specimen  of  C.  altipinnis  (the  specimen  is 
now  the  holotype  of  C.  altipinnis). 

ETYMOLOGY. — From  the  Latin  altus,  high, 
and  pinna,  fin,  in  reference  to  the  unusually  high 
second  dorsal,  anal,  and  first  dorsal  fins. 

Callionymus  sokonumeri  Kamohara,  1936 

Callionymus  sokonumeri  KAMOHARA,  1936:448,  fig.  2  (Mi- 
mase  Market);  1952:90,  fig.  87  (Prov.  Tosa);  1955:63,  fig.; 
MATSUBARA  1955:713  (after  Kamohara);  OCHIAI,  ARAGA, 
AND  NAKAJIMA  1955:109-110,  figs.  6-7,  table  5  (Mimase 
and  Owase);  KAMOHARA  1964:90  (Kochi  Pref.). 

DIAGNOSIS. — A  Callionymus  of  the  kaianus- 
group  with  a  small  branchial  opening;  short  head 
(3.6-4.4  in  SL);  preopercular  spine  with  a  small 
antrorse  and  two  larger  curved  points  at  dorsal 
side,  and  a  short  main  tip;  high  second  dorsal 
fin  with  a  convex  distal  margin;  no  filament  in 
first  dorsal  fin;  anal  fin  with  a  distal  black  streak 
on  each  membrane;  distal  two-thirds  of  mem- 
branes of  anal  fin  brown,  including  tips  of  fin 
rays;  an  irregular  distal  margin  of  caudal  fin;  and 
a  very  small  black  blotch  on  third  membrane  of 
first  dorsal  fin,  extremely  distal  in  position. 

DESCRIPTION. — Counts  and  proportions  (see 
also  Table  2):  D,  IV;  D2  viii.l;  A  viii.l;  P,  18- 
20;  P2  1,5;  C  10. 

Body  elongate  and  depressed.  Head  de- 
pressed, 3.6-4.4  in  SL.  Eye  large,  2.1-2.9  in 
head.  Pupil  about  3.1  in  eye  diameter.  Branchial 
opening  about  3  in  pupil.  Preopercular  spine 
with  a  short  main  tip,  a  small  antrorse  and  two 
larger  curved  points  at  its  dorsal  side,  and  a  large 


antrorse  spine  at  its  base  (formula:  1- 


-1). 


Lateral  line  reaching  from  area  behind  eye  to 
end  of  one  of  the  median  caudal  fin  rays  (?);  the 
line  of  the  opposite  side  interconnected  by  a 
transverse  branch  across  occipital  region  and 


another  across  dorsal  side  of  caudal  peduncle. 
Caudal  peduncle  length  about  4.8  in  SL,  minimal 
caudal  peduncle  depth  23.0  in  SL. 

First  spine  of  first  dorsal  fin  lower  than  first 
ray  of  second  dorsal  fin,  not  filamentous.  Second 
dorsal  fin  very  high,  with  a  convex  distal  margin. 
Anal  fin  begins  on  vertical  through  second  ray 
of  second  dorsal  fin.  Distal  shape  of  caudal  fin 
irregular;  two  median  caudal  fin  rays  elongate, 
occasionally  filamentous.  Distal  margin  of  pelvic 
fin  convex;  longest  pelvic  fin  ray  reaching  to 
midbase  of  first  membrane  of  anal  fin  when  laid 
back.  Pectoral  fin  reaching  to  fourth  ray  of  sec- 
ond dorsal  fin  when  laid  back. 

Color  in  alcohol.  Body  olive-yellow  above, 
whitish  below.  Back  with  both  blackish  spots 
and  blotches,  the  former  forming  darkish  rings. 
A  row  of  several  dark  brown  blotches  along  side 
of  body.  Pectoral  fin  base  with  a  brown  area. 
First  dorsal  fin  gray,  with  three  oblique  dark 
bars;  a  black  blotch  distally  on  third  membrane. 
Second  dorsal  fin  gray,  with  several  vertical 
dark  streaks  on  membranes.  Distal  part  of  anal 
fin  blackish,  occasionally  with  a  transverse 
streak  distally  on  each  membrane  except  the 
first.  Caudal  fin  gray,  mottled  with  blackish 
spots  and  small  whitish  blotches.  Pectoral  fin 
pale,  the  dorsal  part  mottled  with  dark  spots. 
Pelvic  fin  gray,  distal  parts  of  fourth  and  fifth 
rays  dark. 

DISTRIBUTION. — Known  only  from  central 
Honshu,  Japan  (Pacific  coast;  see  Fig.  2);  col- 
lection depths  not  known. 

DISCUSSION. — I  unfortunately  had  no  material 
of  this  species.  The  present  description  is  com- 
piled from  Kamohara  (1936)  and  Ochiai,  Araga, 
and  Nakajima  (1955). 

The  differences  between  C.  sokonumeri  and 
C.  altipinnis  are  discussed  in  the  description  of 
the  latter  species.  Callionymus  sokonumeri  dif- 
fers from  C.  ochiaii  in  its  higher  second  dorsal 
and  anal  fins,  the  shape  of  the  caudal  fin,  a  short- 
er main  tip  of  the  preopercular  spine,  and  var- 
ious color  markings. 

Callionymus  sp. 

Callionymus  kaianus:   (non   Giinther,    1880)   WONGRATANA 
1968:58  (Gulf  of  Thailand). 

DISCUSSION. — I  did  not  examine  material  of 
the  kaianus  -group  from  the  Gulf  of  Thailand. 
Wongratana's  (1968)  paper  records  Callionymus 
kaianus,  but  gives  no  descriptions  or  figures. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  14 


Callionymus  kaianus  should  not  occur  in  the 
Gulf  of  Thailand.  I  have  been  unable  to  deter- 
mine if  the  specimens  from  that  area  belong  to 
any  of  the  known  species  of  the  kaianus 'group, 
or  if  they  represent  yet  another  species. 

ACKNOWLEDGMENTS 

For  discussion  of  problems,  loan  of  speci- 
mens, information,  or  permission  to  examine 
specimens  in  their  collections,  I  thank  the  fol- 
lowing persons:  H.-J.  Paepke  (ZMB,  East  Ber- 
lin); P.  J.  P.  Whitehead,  A.  Wheeler,  and  O. 
Crimmen  (BMNH,  London);  A.  Ben-Tuvia  and 
D.  Golani  (HUJF,  Jerusalem);  K.-C.  Au  (MSL, 
Hong  Kong);  H.  K.  Larson  (AMS,  Sydney);  J. 
P.  Gosse  (IRSN,  Brussels);  W.  N.  Eschmeyer, 
L.  J.  Dempster,  T.  Iwamoto,  T.  R.  Roberts,  P. 
M.  Sonoda,  M.  Hearne,  and  G.  Raabe  (CAS, 
San  Francisco),  T.  Iwai  and  T.  Nakabo  (FAKU, 
Kyoto),  A.  Kotthaus  (Eppstein,  W.  Germany), 
H.  Wilkens  and  R.  Dohse  (ZIM,  Hamburg).  W. 
N.  Eschmeyer  and  T.  Iwamoto  (CAS,  San  Fran- 
cisco) also  reviewed  the  manuscript. 

I  am  greatly  obliged  to  P.  Fourmanoir  (OR- 
STOM,  Noumea)  for  the  gift  of  a  specimen  of 
Callionymus  moretonensis  and  a  further  callion- 
ymid  fish  specimen.  I  am  also  grateful  to  D.  J. 
Stewart  (FMNH,  Chicago)  who  examined  a 
specimen  of  Callionymus  carebares  from  his 
collection  for  me.  I  thank  O.  von  Frisch  (NMB, 
Braunschweig)  for  accepting  specimens  loaned 
to  me.  A  financial  contribution  for  expenses  dur- 
ing my  visit  to  the  California  Academy  of  Sci- 
ences was  provided  by  that  organization. 

LITERATURE  CITED 

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MATSUBARA,  K.  1955.  Fish  morphology,  and  hierarchy.  Pt. 
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MUNRO,  I.  S.  R.  1958.  The  fishes  of  the  New  Guinea  region. 
Territ.  Papua  and  New  Guinea,  Fish.  Bull.  1:97-369.  [Re- 
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369.] 

NAKABO,  T.  1979.  A  new  and  two  rare  species  of  the  genus 
Callionvmus  (Callionymidae)  from  the  western  Indian 
Ocean.  Japan.  J.  Ichthyol.  26(3):231-237,  figs,  l^t,  table  1. 


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132. 

PETERS,  W.  C.  H.  1876.  Uebersicht  der  wahrend  der  von 
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CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  15,  pp.  379^407,  18  figs.,  4  tables. 


October  26,  1981 


DISTRIBUTION,  REPRODUCTIVE  ANATOMY,  AND 

VARIATION  OF  MONADENIA  TROGLODYTES  HANNA 

AND  SMITH  (GASTROPODA:  PULMONATA)  WITH 

THE  PROPOSAL  OF  A  NEW  SUBGENUS 

By 
Barry  Roth 

Department  of  Invertebrate  Zoology,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  94118 


ABSTRACT:  Shastelix,  new  subgenus  of  Monadenia  (type-species,  M.  troglodytes  troglodytes)  is  proposed, 
based  principally  on  genital  characters.  The  new  subgenus  is  mostly  confined  to  the  Klamath  Mountains  of 
California  and  is  parapatric  with  Monadenia,  sensu  stricto.  Monadenia  troglodytes  is  common  in  a  limited  area 
of  Shasta  County,  associated  with  limestone  terrane.  Three  subspecies  are  recognized  based  on  morphometry 
and  geography — M.  t.  troglodytes  along  the  McCloud  River  arm  of  Shasta  Lake;  M.  t.  wintu,  new  subspecies, 
between  the  Pit  River  and  Squaw  Creek,  with  one  disjunct,  outlying  population  south  of  Shasta  Lake;  and  M. 
t.  chaceana  (formerly  ranked  as  a  distinct  species)  near  the  confluence  of  the  Shasta  and  Klamath  rivers  in 
Siskiyou  County.  Sympatric  Monadenia  churchi  and  M.  troglodytes  differ  little  in  reproductive  anatomy;  elab- 
orations of  the  female  genitalia  are  probably  not  important  in  species  recognition  or  reproductive  isolation.  A 
phylogenetic  hypothesis  and  evolutionary  scenario  for  the  three  subgenera  of  Monadenia  are  presented. 


INTRODUCTION 

The  western  North  American  helicacean  snail 
genus  Monadenia  Pilsbry,  1895,  includes  con- 
spicuous species  that  have  long  attracted  the 
attention  of  malacologists.  Its  type-species,  He- 
lix fidelis  Gray,  1834,  was  the  first  land  mollusk 
described  from  the  Pacific  coast.  It  includes  the 
most  northern  helicacean  species — and  some  of 
the  most  northern  large  land  snails — on  the 
North  American  continent.  Monadenia  is  one 
of  the  few  temperate  genera  of  snails  with  an 
elaborate  color  pattern  (Comfort  1951),  and  M. 
fidelis  is  the  only  snail  in  the  far  west  with  a 
dramatic  polymorphism  of  shell  color  and  band- 
ing (Roth  in  press). 

Pilsbry  (1939)  and  Berry  (1940a)  divided  Mon- 
adenia into  two  groups  of  species,  based  on  re- 


productive anatomy  and  shell  characters — Mon- 
adenia, sensu  stricto,  and  Corynadenia  Berry, 
1940.  The  range  of  the  genus  extends  from 
southern  Alaska  to  central  California,  principal- 
ly west  of  the  Cascade  Range  but  penetrating 
inland  along  major  river  valleys.  One  branch 
reaches  south  along  the  east  side  of  the  Sacra- 
mento Valley;  the  other  follows  the  southern 
Klamath  Mountains  and  Coast  Ranges  to  the 
San  Francisco  Bay  region  (Figure  1).  The  sub- 
genus  Corynadenia  exists  as  an  apparently  dis- 
junct group  of  species  on  the  west  side  of  the 
Sierra  Nevada.  Roth  (1975)  showed  that  Mon- 
adenia churchi  Hanna  and  Smith,  1933,  which 
occupies  a  fairly  wide  range  around  the  north 
end  of  the  Sacramento  Valley,  and  which  Pilsbry 
(1939)  and  Berry  (1940a)  had  grouped  with  the 


[379] 


380 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


FIGURE  1.  Distribution  of  the  subgenera  of  Monadenia  in 
California.  Diagonal  hachure,  generalized  range  of  Moniulen- 
ici,  sensu  stricto;  stippling,  in  Klamath  Mountains  region, 
Shastelix,  in  Sierra  Nevada,  Corynadenia.  Dashed  line  rep- 
resents 36  C  mean  maximum  July  isotherm;  dot-dashed  line, 
-4  C  mean  minimum  January  isotherm  (thermal  data  after 
Elford  1970). 


Sierran  subgenus,  combines  anatomic  and  shell 
characters  of  Monadenia,  s.s.,  and  Corynaden- 
ia. A  number  of  other  species  from  northern 
California  described  by  Berry  (1940b)  have  not 
yet  been  dissected  or  assigned  unequivocally  to 
subgenus.  Parapatry  and  possible  sympatry  of 
species  have  been  demonstrated  (Roth  and  Eng 
1980;  and  herein),  and  it  is  clear  that  the  zoo- 
geography of  Monadenia  is  more  complex  than 
earlier  authors  imagined. 

Monadenia  is  ecologically  diverse,  compris- 
ing species  of  rock  crevices  and  rockslides,  oth- 
ers that  inhabit  deep  leaf  mold,  semi-arboreal 
forms,  and  snails  that  crawl  out  exposed  on  low 
plant  cover.  The  relations  between  habit  and 
habitat,  on  the  one  hand,  and  morphology  and 
variation,  on  the  other,  are  important  for  under- 
standing the  evolutionary  history  of  the  group. 
This  paper  is  one  of  a  projected  series  of  studies 
aimed  at  elucidating  those  relationships. 

Monadenia  troglodytes   Hanna  and  Smith, 


1933,  was  described  from  fossil  shells  of  pre- 
sumed Pleistocene  age  collected  in  Samwel 
Cave,  Shasta  County,  California  (sec.  5,  T.  35 
N,  R.  3  W,  Mount  Diablo  Base  and  Meridian, 
USGS  Bollibokka  Mountain  Quadrangle).  Other 
shells  of  the  species,  likewise  interpreted  as 
Pleistocene,  were  reported  from  Potter  Creek 
Cave  (sec.  23,  T.  34  N,  R.  4  W,  MDB&M, 
USGS  Lamoine  Quadrangle),  Shasta  County 
(Hanna  and  Smith  1933).  Empty  shells  were  lat- 
er found  by  Stanford  University  speleologists  a 
short  distance  outside  Samwel  Cave  (Smith 
1957),  and  in  1963  the  species  was  discovered 
alive  in  the  same  general  area  (Walton  1970). 
Smith  (1970)  and  Roth  (1972a,  1972b)  cited  M. 
troglodytes  as  rare  and  of  limited  distribution. 

In  May  1973  the  late  Allyn  G.  Smith  of  the 
California  Academy  of  Sciences  found  the 
species  at  a  new  locality,  in  limestone  rockslides 
near  Ellery  Creek  (SEV4  sec.  6,  T.  35  N,  R.  3 
W,  MDB&M,  Bollibokka  Mountain  Quadran- 
gle), west  of  the  McCloud  River  arm  of  Shasta 
Lake  (Fig.  2).  He  collected  many  empty  shells 
in  various  states  of  preservation  and  one  living 
specimen.  Since  then,  additional  collections,  in- 
cluding a  substantial  amount  of  material  secured 
independently  by  S.  E.  Hirschfeld,  D.  C.  Ru- 
dolph, and  R.  L.  Seib,  indicate  that  the  species 
is  fairly  common  in  a  limited  area  in  Shasta 
County  and  is  strongly  associated  with  lime- 
stone terrane. 

Dissections  of  the  reproductive  system  show 
that  M.  troglodytes,  along  with  M.  churchi,  be- 
longs to  a  new  subgenus,  which  is  named  herein. 
Selected  shell  characters  were  measured  to  ana- 
lyze shell  variability.  A  new  subspecies  is  de- 
scribed, distinguished  from  typical  M.  troglo- 
dytes by  details  of  color,  shell  microsculpture, 
and  morphometry.  Monadenia  chaceana  Berry 
(1940b)  is  similar  in  general  shell  character  but 
differs  consistently  in  certain  shell  measure- 
ments and  is  regarded  as  a  third  subspecies.  One 
other  species  of  Monadenia  occurs  within  the 
range  of  M.  troglodytes,  permitting  a  consider- 
ation of  species  criteria  within  the  genus.  An 
hypothesis  of  phylogenetic  relationships  within 
Monadenia  is  presented. 

The  use  of  a  trinomial  to  designate  the  nomi- 
nate subspecies — a  convention  sparingly  ob- 
served in  American  land  malacology,  but  one 
necessary  to  distinguish  the  subspecies  from  the 
species  sensu  lato — is  here  introduced  for  Mon- 
adenia troglodytes  troglodytes. 


ROTH:  DISTRIBUTION,  ANATOMY,  VARIATION  OF  MONADENIA  TROGLODYTES 


381 


FIGURE  2.     Oilman  Road  crossing  of  Ellery  Creek  (Locality  26);  typical  Monadenia  troglodytes  habitat  in  brush-covered 
talus  at  foot  of  prominent  limestone  outcrop. 


The  following  institutional  abbreviations  are 
employed: 

AMNH — American  Museum  of  Natural  History 

ANSP — Academy  of  Natural  Sciences,  Phila- 
delphia 

CAS — California  Academy  of  Sciences,  Depart- 
ment of  Invertebrate  Zoology 

CASGTC — California  Academy  of  Sciences, 
Geology  Type  Collection,  Department  of  Ge- 
ology 

FMNH — Field  Museum  of  Natural  History 

LACM — Los  Angeles  County  Museum  of  Nat- 
ural History 

SSB — Private  collection  of  S.  Stillman  Berry, 
Redlands,  California 

SUPTC— Stanford  University  Paleontological 
Type  Collection,  now  in  Department  of  Ge- 
ology, California  Academy  of  Sciences 

UCMP — Museum  of  Paleontology,  University 
of  California,  Berkeley 

USNM — United  States  National  Museum  of 
Natural  History 


Monadenia  Pilsbry 

Monadenia  PILSBRY,  1895:198.— PILSBRY  1939:31-35. 
TYPE-SPECIES:  Helix  fidelis  Gray,  1834,  by  original  designa- 
tion. 

Shastelix,  new  subgenus 

TYPE-SPECIES:  Monadenia  troglodytes  troglodytes  Hanna  and 
Smith,  1933. 

DIAGNOSIS. — Monadenia  with  large,  globose 
atrium;  mucus  gland  much  longer  than  dart  sac, 
its  lower  part  adnate  to  atrium;  penial  retractor 
inserted  near  middle  of  epiphallus;  flagellum 
(epiphallic  caecum)  substantially  longer  than  pe- 
nis plus  epiphallus  and  borne  in  a  series  of  he- 
lical coils.  Penis  sessile  on  atrium,  not  invagi- 
nated  into  it.  Spermatophore  helically  coiled 
(?).  Shell  of  moderate  size  for  the  genus,  smooth 
or  granulose,  protoconch  sculpture  of  minute, 
somewhat  confluent  granules,  tending  to  align  in 
diagonal  series. 

The  genitalia  of  Monadenia  troglodytes  (Figs. 
4-6)  differ  most  obviously  from  those  of  species 


382 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


TABLE  1.     CHARACTER  COMPLEXES  IN  THE  SUBGENERA  OF  Monadenia. 


Characters 


Monadenia,  s.s. 


Shastelix 


Corvnadenia 


Atrium  large  and  globose  (+),  or  small  and  narrow 
(-) 

Mucus  gland  many  times  longer  than  (  +  ),  or  about 
equal  to  (-),  dart  sac 

Lower  part  of  mucus  gland  adnate  to  (  +  ),  or  free  from 
(-),  atrium 

Dart  sac  5  mm  or  less  in  length  (+),  or  longer  than 
5  mm  (-) 

Flagellum  ^\.5  times  length  of  (+),  or  about  as  long 
as  (-),  penis  plus  epiphallus 

Flagellum  (and  spermatophore)  helically  coiled  (+), 
or  straight  (-) 

Basal  chamber  of  penis  invaginated  into  (+),  or 
sessile  on  (-),  atrium 

Penial  retractor  inserted  near  middle  (+),  or  on 
distal  third  (-),  of  epiphallus 


of  the  nominate  subgenus  (M.fidelis,  M.  infu- 
mata  (Gould,  1855),  M.  setosa  Talmadge,  1952) 
in  the  long,  helically  coiled  flagellum.  The  fla- 
gellum  in  Monadenia,  sensu  stricto,  is  thick  and 
straight  or  simply  curved  and  about  as  long  as 
the  penis  plus  epiphallus  (Pilsbry  1939:figs.  15A, 
B,  M.fidelis;  Roth  and  Eng  1980:fig.  3,  M.  se- 
tosa; the  genitalia  of  M.  infumata  are  similar). 
The  flagellum  is  the  organ  which  secretes  the 
spermatophore;  the  spermatophore  of  M.fidelis 
(Webb  1952:fig.  8A)  is  straight.  In  the  Sierran 
Monadenia  (Corvnadenia)  hirsuta  Pilsbry, 
1927,  a  species  with  helically  coiled  flagellum, 
the  spermatophore  is  coiled  like  a  corkscrew.  It 
seems  probable,  therefore,  that  species  of  Shas- 
telix also  secrete  coiled  spermatophores. 

Both  Shastelix  and  Monadenia,  s.s.,  have  a 
large,  globose  atrium.  (I  follow  Pilsbry  [1939], 
Berry  [1940a]  and  other  authors  in  using  the 
term  "atrium"  for  the  large,  saccular  elabora- 
tion of  the  lower  genitalia  upon  which  the  penis, 
dart  sac,  and  vagina  insert  and  which  when 
everted  forms  the  copulatory  pad  or  disk.  This 
organ  is  mainly  developed  above  the  insertion 
of  the  penis  and  is  homologous  to  the  lower  part 
of  the  vagina  in  other  helicoid  genera.  In  these 
other  genera  the  term  "atrium"  is  convention- 
ally restricted  to  the  common  passage  to  the  ex- 
ternal genital  pore  below  the  insertion  of  the  pe- 
nis.) In  Shastelix  the  lower  ductlike  portion  of 


the  mucus  gland  is  adnate  to  the  atrium,  whereas 
in  Monadenia,  s.s.,  it  runs  along  the  surface  of 
the  atrium  but  is  not  fused  to  it.  The  dart  sac  in 
Shastelix  is  smaller  than  that  in  the  nominate 
subgenus.  In  Monadenia,  s.s.,  the  basal  part  of 
the  penis  is  invaginated  into  the  wall  of  the 
atrium,  which  clasps  it  like  a  collar;  in  Shastelix 
the  basal  part  of  the  penis  is  sessile  on  the 
atrium. 

In  the  subgenus  Corvnadenia  Berry,  1940a 
(type-species,  Helix  hillebrandi  Newcomb, 
1864;  see  Pilsbry  1939:fig.  15C),  the  atrium  is 
smaller,  narrow  and  elongate,  and  the  mucus 
gland  is  shorter  or  very  slightly  longer  than  the 
dart  sac.  The  flagellum  is  longer  than  the  penis 
plus  epiphallus  and,  at  least  in  some  species, 
helically  coiled.  The  basal  part  of  the  penis  is 
invaginated  into  the  wall  of  the  atrium,  as  in 
Monadenia,  s.s. 

Character  complexes  differentiating  the  three 
subgenera  are  summarized  in  Table  1. 

Monadenia  churchi  also  belongs  to  Shastelix 
and  resembles  M.  troglodytes  in  genitalia  (Fig. 
8)  and  protoconch  sculpture.  It  is  probable  that 
some  undissected  species  from  the  Klamath 
Mountains  (particularly  M.  cristulata  Berry, 
1940,  and  M.  marmarotis  Berry,  1940)  will  also 
prove  to  belong  to  Shastelix.  The  known  range 
of  the  subgenus  (Fig.  1)  extends  from  Butte 
County  on  the  south  and  east,  around  the  north 


ROTH:   DISTRIBUTION.  ANATOMY.  VARIATION  OF  MONADEN1A  TROGLODYTES 


383 


^•S-XV         R3W 

'8,9 
[JlilO-24 

25< 


R2W 


130 


35 


4Q.41 


Monadenia  churchi 

M.  troglodytes  troglodytes 

M.  troglodytes  wintu 

M.  churchi  and  M.  troglodytes 


FIGURE  3.  Distribution  of  Monadenia  species  in  vicinity  of  Shasta  Lake,  Shasta  County,  Calif.  Stippling  indicates  surface 
outcrops  of  limestone.  Open  triangles  denote  peaks  of  more  than  1000  m  elevation.  Geology  after:  Diller  (1906);  Kinkel  et  al. 
(1956);  Coogan  (1960);  Albers  and  Robertson  (1961);  Demirmen  and  Harbaugh  (1965);  Irwin  and  Galanis  (1976). 


end  of  the  Sacramento  Valley,  to  near  Burnt 
Ranch,  Trinity  County,  on  the  west,  and  the 
Shasta  River-Klamath  River  confluence  on  the 
north.  Monadenia  churchi  and  M.  troglodytes 
are  sympatric  in  the  vicinity  of  Shasta  Lake.  In 
the  drainage  of  the  Trinity  River,  Shastelix  is 
parapatric  with  Monadenia,  s.s.  There  M.  fi- 
delis  and  M.  setosa  tend  to  occur  in  riparian 
woodland  close  to  streambeds,  while  M.  churchi 
occurs  on  drier,  more  exposed  slopes  and  in 
rockslides.  Only  the  subgenus  Corynadenia  is 
known  to  occur  in  the  Sierra  Nevada. 

The  subgenus  name  is  derived  from  the  place- 
name  Shasta — lake,  mountain,  river,  and 
county — plus  the  Greek  helix,  a  spiral,  hence  a 
snail.  Its  gender  is  feminine. 


DISTRIBUTION,  ANATOMY,  AND  GEOGRAPHIC 
VARIATION  OF  Monadenia  troglodytes 

The  material  now  available,  summarized  in 
the  Appendix,  makes  it  clear  that  Monadenia 
troglodytes  is  not  only  extant  (Walton  1970)  but 
occupies  a  considerable  range  in  the  vicinity  of 
Shasta  Lake  (Fig.  3).  At  several  of  the  localities, 
living  individuals  were  found  to  be  common. 

The  ranges  of  M.  t.  troglodytes  and  a  second 
subspecies  described  herein  lie  within  the 
"Eastern  Klamath  Belt/'  the  easternmost  of 
several  concentric  lithic  belts  that  constitute  the 
Klamath  Mountains  physiographic  province  of 
northwest  California  and  southwest  Oregon  (Ir- 
win 1972).  The  Klamath  Mountains  are  a  rugged 


384 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42.  No.  15 


mountain  region  including  clusters  of  high  peaks 
1800-2700  m  in  altitude.  Peaks  and  ridgecrests 
of  1200-1350  m  occur  in  the  area  mapped  in  Fig- 
ure 3.  The  canyons  of  the  Sacramento,  Mc- 
Cloud,  and  Pit  rivers  and  Squaw  Creek  dominate 
the  area's  topography.  Shasta  Lake  is  an  artifi- 
cial impoundment  of  these  rivers;  their  courses 
prior  to  damming  may  be  seen  on  pre-1940  to- 
pographic maps.  The  spillway  of  Shasta  Dam  is 
at  an  elevation  of  325  m,  so  that  local  relief  of 
1000  m  and  more  is  present  in  the  area  under 
study.  Southward  from  Shasta  Lake,  the  Sac- 
ramento River  descends  through  foothills  to  the 
broad,  alluviated  floor  of  the  Sacramento  Val- 
ley. The  Sacramento  River-Trinity  River  wa- 
tershed— closely  coinciding  with  the  Shasta 
County-Trinity  County  boundary  and  dividing 
the  north  coast  drainage  from  the  interior,  Great 
Valley,  drainage — is  located  a  few  kilometers  off 
the  west  margin  of  the  map.  North,  east,  and 
southeast  of  the  map  area  lie  portions  of  the 
Cascade  Range  physiographic  province  (Wahr- 
haftig  and  Birman  1965);  due  north  is  the  Pleis- 
tocene stratovolcano  Mount  Shasta,  4300  m  in 
altitude. 

The  Klamath  Mountains  are  noted  for  the  di- 
versity of  their  flora  and  the  number  of  local 
endemic  plant  species  (Whittaker  1961;  Axelrod 
1976).  In  the  region  under  study,  the  vegetation 
is  largely  northern  yellow  pine  forest  with  some 
Sierran  montane  forest  (Kuchler  1977),  partic- 
ularly between  the  Pit  and  McCloud  rivers. 
Lower  elevations  and  exposed  slopes  are  cov- 
ered by  a  growth  of  scrub  oak  and  chaparral. 
The  land  at  higher  elevations  was  originally 
densely  timbered  with  coniferous  forest,  but  in 
many  areas  logging,  fires,  and  the  effects  of 
smelter  smoke  have  removed  the  original  tim- 
ber, so  that  second-growth  forest  and  brush  are 
widespread.  Manzanita  (Arctostaphylos)  is  the 
dominant  brush  plant. 

The  normal  annual  precipitation  is  between 
120  and  180  cm,  more  than  90  percent  of  it  falling 
in  the  months  October  through  April  (Elford 
1970;  Major  1977).  Mean  monthly  temperatures 
at  Shasta  Dam  range  from  around  5  C  in  January 
to  about  25  C  in  July  (Major  1977).  Daytime  tem- 
peratures in  excess  of  40  C  are  not  uncommon 
from  June  through  September.  Local  microcli- 
matic  variation  is  considerable. 

Geology  of  the  region  is  complex,  including 
intrusive  and  extrusive,  sedimentary,  and  meta- 


morphic  rock  suites,  of  ages  from  Paleozoic  to 
Quaternary.  Of  particular  interest  are  bodies  of 
limestone  that  crop  out  discontinuously  in  three 
principal  areas.  These  are  finely  crystalline,  re- 
sistant limestones  that  often  form  prominent 
outcrops  (Kinkel  et  al.  1956:fig.  20).  Local  to- 
pographic names  such  as  "Gray  Rocks"  (La- 
moine  and  Bollibokka  Mountain  quadrangles), 
"Limerock  Gulch,"  "Marble  Creek,"  and  "Up- 
per and  Lower  Limestone  Valley  Creeks"  (La- 
moine  Quadrangle)  acknowledge  their  presence. 

All  samples  of  Monadenia  troglodytes  for 
which  adequate  data  are  available  were  taken  on 
or  adjacent  to  these  limestone  areas  (Fig.  3).  The 
localities  fall  into  two  groups:  those  associated 
with  the  Triassic  Hosselkus  limestone  in  the  vi- 
cinity of  Brock  Mountain,  between  the  Pit  River 
and  Squaw  Creek  arms  of  Shasta  Lake;  and 
those  on  or  near  the  Permian  McCloud  lime- 
stone, along  the  McCloud  River  arm  of  the 
Lake.  The  type-locality,  Samwel  Cave,  belongs 
to  the  latter  group;  so  does  Locality  45,  south 
of  the  Pit  River  arm,  the  southernmost  definite 
site  for  M.  troglodytes.  (A  bleached,  broken 
shell  from  somewhat  farther  south — Loc.  46,  in 
the  Anderson  Quadrangle — is  similar  but  cannot 
be  assigned  unequivocally  to  species.)  Locality 
35  is  from  a  limestone  quarry,  evidently  in  one 
of  the  limestone  lenses  in  the  predominantly 
clastic  Pit  Formation  (Albers  and  Robertson 
1961). 

Limestones  in  the  Shasta  Lake  region  are  dis- 
continuous because  of  faulting  and  erosion.  The 
McCloud  and  Brock  Mountain  locality  groups 
are  separated  by  approximately  10  km  of  non- 
limestone  terrane — chiefly  volcanics,  pyroclas- 
tics,  and  mudstones.  Squaw  Creek,  its  canyon, 
and  a  watershed  of  830  m  minimum  elevation 
also  stand  between  the  two  groups.  Morphologic 
distinctions  between  the  Brock  Mountain  and 
McCloud  River  herds,  discussed  below,  indicate 
that  a  certain  amount  of  genetic  isolation  exists 
as  well.  Localities  yielding  M.  troglodytes  range 
in  elevation  from  about  330  to  760  m,  so  it  does 
not  seem  likely  that  the  altitude  of  the  watershed 
alone  constitutes  an  isolating  factor.  Exposure 
and  seasonal  lack  of  moisture  on  ridgetops  may 
restrict  the  snails1  mobility,  but  since  some  col- 
lections of  M.  troglodytes  were  made  in  zones 
of  high  insolation,  with  snails  active  in  shaded 
spots  even  during  July,  limestone  substrate 
seems  a  more  likely  limiting  factor. 


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385 


Cooper  (1869)  remarked  on  the  strong  asso- 
ciation of  Monadenia  with  carbonate  terrane  in 
the  Sierra  Nevada;  and  Pilsbry  (1939)  surmised 
that  discontinuity  of  limestone  substrata  was  in- 
volved in  the  formation  of  local  races.  Many 
other  records  exist  of  land  mollusks  whose  dis- 
tribution is  more  or  less  tied  to  areas  of  high 
calcium  availability  (e.g.,  Boycott  1934;  Twee- 
die  1961;  Heller  1975). 

To  the  west,  limestone  outcrops  of  the  De- 
vonian Kennett  Formation  in  the  Mammoth 
Buttes-Backbone  Ridge  area  west  of  the  Sac- 
ramento River  arm  of  Shasta  Lake  have  yielded 
Monadenia  churchi  but  not  M.  troglodytes. 
Monadenia  churchi  also  occurs  in  the  McCloud 
River  and  Brock  Mountain  areas.  At  a  minimum 
of  three  stations  it  is  sympatric  or  parapatric 
with  M.  troglodytes  and  was  received  in  the 
same  samples.  M.  churchi  is  not  restricted  to 
limestone  substrata;  it  has  been  found  elsewhere 
in  lava  rockslides  (type  lot;  Hanna  and  Smith 
1933)  and  coniferous  forest  debris  (Roth  and  Eng 
1980).  At  each  locality  on  limestone  where  both 
M .  troglodytes  and  M .  churchi  were  taken  to- 
gether, M.  troglodytes  is  the  more  common 
species.  The  implications  of  the  sympatry  of  M. 
churchi  and  M.  troglodytes  for  classification  of 
Monadenia  are  discussed  below. 

No  specimens  from  Shasta  County  referable 
to  the  Sierran  Monadenia  (Corynadenia)  mor- 
monum  (Pfeiffer,  1857)  have  been  found  in  the 
course  of  this  study,  and  it  appears  that  all  such 
records  in  the  literature  are  based  on  misiden- 
tifications  of  either  M.  troglodytes  or  M.  chur- 
chi. (For  example:  "The  most  northern  locality 
for  mormonum  now  known  is  at  Shasta,  Cal., 
lat.  41°  (nearly),  alt.  1160  feet,  where  in  the  vol- 
canic region  Dr.  Yates  found  a  very  few  stunted 
specimens  with  but  five  and  a  half  whorls  and 
the  bristle-granulations  of  the  young  very 
strongly  developed"  [Cooper  1879:285].  "In 
Shasta  County,  far  north  of  the  localities  men- 
tioned, a  race  of  mormonum  has  been  found  in 
the  Upper  Sonoran  Zone  at  and  near  the  junc- 
tion of  the  Pitt  [sic]  with  the  Sacramento  river 
(Brewer,  Gabb).  They  agree  with  the  typical 
form  in  the  absence  or  extreme  faintness  of  spi- 
ral striae  and  in  coloration;  the  shell  is  smaller 
and  the  spire  generally  higher.  They  are  within 
the  area  of  M.  churchi"  [Pilsbry  1939:56].)  A  lot 
collected  in  the  nineteenth  century,  UCMP 
2491,  ex  D.  O.  Mills  collection  no.  290,  labeled 


"Shasta  County,  Calif."  and  formerly  identified 
as  M.  mormonum,  is  probably  the  same  as  the 
Shasta  County  shells  referred  to  by  Pilsbry 
(1939:56).  It  is  M.  troglodytes,  similar  to  those 
from  the  Brock  Mountain  area.  The  original  de- 
scription of  M.  troglodytes  was  based  on  sam- 
ples with  very  low  spire  index  (H/D  =  0.411- 
0.500);  the  figured  holotype  is  nearly  planispiral. 
In  the  samples  now  at  hand,  intermediate  ex- 
amples connect  these  very  flat  shells  with  the 
higher-spired,  mormonum-\ike  specimens. 

ANATOMY. — Specimens  were  prepared  for 
anatomical  study  by  drowning  followed  by 
transfer  in  stages  to  70%  ethanol.  Specimens  of 
M.  troglodytes  from  the  following  localities 
were  dissected:  10,  26,  28,  34,  39,  43.  Except  as 
noted,  the  data  given  apply  to  all  individuals  ex- 
amined. 

Body  grayish  tan,  shaded  darker  on  dorsum; 
light  buff  mid-dorsal  stripe;  sole  light  buff.  Man- 
tle over  lung  translucent  buff,  with  gray  dendri- 
tic pencilling  occupying  0-35%  (usually  about 
20%)  of  surface.  (All  degrees  of  mantle  pigmen- 
tation present  in  sample  from  Loc.  10;  other 
samples  less  variable.)  Mantle  collar  colored  like 
body.  External  genital  pore  a  vertical  1-2  mm 
slit  often  showing  white  rim. 

Right  ocular  retractor  running  between  male 
and  female  systems,  passing  over  crook  at  penis- 
epiphallus  junction.  Genitalia  as  in  Figures  4-6. 
Penis  stout,  basal  chamber  separated  by  crook 
from  upper  chamber,  adnate  to  atrium  but  not 
invaginated  therein;  upper  chamber  thin- walled, 
finely  ridged  internally,  containing  cylindrical  to 
ovate-conic,  slightly  rugulose  verge  3-3.5  mm 
long.  Tip  of  verge  with  slitlike  lateral  meatus  on 
anterior  edge,  dorsal  facet  concavely  beveled 
(Fig.  4b).  (End  of  verge  of  specimens  from  Loc. 
43  blunt,  as  broad  or  broader  than  stalk,  with 
flaplike  expansion  of  tip  opposite  meatus.)  Wall 
of  upper  penial  chamber  bearing  a  single  large 
pilaster  that  fits  against  beveled  facet  of  verge 
and  extends  into  basal  chamber.  Epiphallus 
from  l/2  to  %  as  thick  as  penis  at  their  junction. 
Penial  retractor  (originating  on  floor  of  lung) 
long,  narrow,  slightly  expanded  just  before  in- 
sertion on  medial  part  of  epiphallus.  Flagellum 
(epiphallic  caecum)  longer  than  penis  plus  epi- 
phallus, as  thick  as  epiphallus,  borne  in  4  to  7 
helical  coils;  distal  end  tapering  to  fine  point. 
Vas  deferens  with  2-3  convolutions  where  it 
passes  under  atrium.  Spermatheca  (bursa  cop- 


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FIGURES  4-5.  Genitalia  of  Monadenia  troglodytes.  Figs.  4a-b.  M.  t.  troglodytes;  (a.)  dorsal  aspect,  Loc.  28,  atrium  rotated 
90°  downward  with  respect  to  oviduct;  (b.)  detail  of  verge,  Loc.  10.  Figs.  5a-b.  M.  t.  wintu,  n.  subsp.,  holotype  CAS  018431, 
Loc.  34;  (a.)  dorsal  aspect;  (b.)  ventral  aspect.  Abbreviations:  a — atrium;  ag — albumen  gland;  ds — dart  sac;  ep — epiphallus; 
/? — flagellum;  fo — free  oviduct;  gp — external  genital  pore;  hcl — hermaphroditic  duct;  ing — mucus  gland;  ph — basal  chamber  of 
penis;  pr — penial  retractor  muscle;  pu — upper  chamber  of  penis;  sp — spermatheca:  spd — duct  of  spermatheca;  t — talon  (fer- 
tilization pouch);  vd — vas  deferens;  ve — verge. 


ulatrix)  globose;  spermathecal  duct  long,  straight, 
unbranched,  capacious  for  lower  three-fourths, 
narrower  below  spermatheca.  Dart  sac  small  to 
moderate-sized,  sessile  at  convergence  of  two 
atrial  crura  which,  when  everted,  form  a  copu- 
latory  pad  or  disk  (Fig.  7);  sac  containing  a 
2-mm,  tubular,  calcareous  dart.  Mucus  gland 
large,  bent  near  middle,  lower  portion  adnate  to 


atrium  and  running  along  lower  edge  of  superior 
atrial  crus  to  insertion  at  base  of  dart  sac. 

Measurements  of  selected  organs  are  given  in 
Table  2.  All  show  considerable  variation.  There 
appears  to  be  no  systematic  difference  between 
samples  except  that  those  from  the  Brock 
Mountain  area  tend  to  have  more  coils  to  the 
epiphallus. 


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387 


TABLE  2.     LENGTHS  (in  mm)  OF  SELECTED  ORGANS  OF  Monadenia  troglodytes  AND  M.  churchi  (range,  with  sample  mean 
in  parentheses). 


Loc.  (n) 

M.  t. 

troglodytes 

M.  t.  wintu 

M.  churchi 

10(6) 

43(5) 

34  (3)                 45  (  1) 

Mammoth 
Butte  (3) 

29(1) 

Free  portion  of  mucus  gland 

16.0-21.2 
(18.82) 

10.9-21.0 
(17.12) 

16.1-30.0               15.7 
(21.0) 

15.7-18.1 
(16.77) 

17.3 

Dart  sac 

2.6-4.5 
(3.18) 

2.1-2.4 
(2.33) 

2.5-3.2                  2.1 
(2.87) 

1.8-2.7 
(2.23) 

1.9 

No.  of  coils  of  flagellum 

3.5-5 
(4.13) 

3.5-5 
(4) 

5-7                     5 
(6) 

3.5-4 
(3.67) 

6 

Free  (upper)  chamber  of  penis 

3.3-4.8 
(3.55) 

3.4-4.7 
(3.84) 

3.7^.6                  4.0 
(4.23) 

3.5-3.6 
(3.6) 

3.5 

Epiphallus 

5.6-8.7 
(7.93) 

5.7-8.6 

(6.72) 

8.4-8.9                  8.1 
(8.63) 

5.2-6.5 
(5.63) 

6.6 

Base  of  epiphallus  to  insertion 
of  penial  retractor 

2.6-4.9 
(4.07) 

2.9-4.1 
(3.5) 

3.5_4.7                  4.1 
(4.03) 

2.2-3.2 
(2.7) 

2.6 

In  their  exserted  state  the  atrial  crura  form  a 
copulatory  pad  (Fig.  7).  The  female  genital  pore 
is  located  on  the  bottom  of  the  pocket  between 
the  two  crura.  The  male  opening,  through  which 
the  verge  is  exserted,  is  located  near  the  middle 
of  the  forward  edge  of  the  anterior  crus.  The 
pore  of  the  dart  sac  is  at  the  convergence  of  the 
crura;  further  eversion  of  the  dart  apparatus  was 
not  observed. 

The  lower  genitalia  are  enveloped  in  thin  tis- 
sue that  binds  the  male  system  to  the  atrium  and 
in  some  specimens  forms  a  collar  around  the 
penis  just  below  its  junction  with  the  epiphallus. 

The  jaw  is  crescentic,  golden  brown,  with  four 
to  six  major  ribs  that  denticulate  both  margins. 
From  one  to  five  smaller  intercalary  ribs  are 
present  on  some  specimens. 

The  radula  of  a  specimen  from  Locality  28  has 
41  teeth  in  a  half  row.  The  central  tooth  is  some- 
what wider  than  in  M.  fidelis  (see  Pilsbry 
1939:fig.  15E),  the  mesocone  as  long  as  the  basal 
plate.  On  the  fourteenth  tooth  an  endocone  ap- 
pears, and  on  the  fifteenth  an  ectocone.  The 
marginal  teeth  are  somewhat  spatulate,  the  cleft 
between  mesocone  and  endocone  deep,  and 
both  mesocone  and  endocone  are  sometimes 
bifid  or  trifid.  The  outer  marginals  are  very 
short. 

SHELL  CHARACTERS  AND  VARIATION. — Han- 
na  and  Smith  (1933)  described  the  protoconch 
of  Monadenia  troglodytes  as  consisting  of  \l/2 
whorls  marked  by  wavy  radiating  riblets.  In  a 


group  of  19  M.  t.  troglodytes  hatched  in  captiv- 
ity from  eggs  laid  by  an  individual  from  Locality 
10  (slope  below  Samwel  Cave),  the  protoconch 
consists  of  1.5-1.8  whorls  (counting  by  the 
method  of  Diver  1932)  and  measures  2.8-3.5  mm 
in  diameter.  The  extreme  nuclear  tip  is  smooth. 
Thereafter,  sculpture  of  the  protoconch  consists 
of  minute  granules,  round  and  somewhat  con- 
fluent, irregularly  disposed  at  first  but  after 
about  one  half  whorl  tending  to  form  wavy,  ra- 
dial riblets.  Those  just  outboard  of  the  suture 
retain  this  character  nearly  to  the  end  of  the  em- 
bryonic shell.  Elsewhere  on  the  top  surface  the 
granules  become  discrete  and  spirally  elongated 
and  tend  to  align  in  oblique,  protractive  and  re- 
tractive series.  Below  the  periphery  the  granules 
are  discrete,  spirally  elongated,  sometimes  ar- 
ranged in  diagonal  series  but  more  often  scat- 
tered. All  other  M.  troglodytes  in  which  the  pro- 
toconchs  are  not  abraded  show  essentially  the 
same  sculpture.  Slight  erosion  sometimes  em- 
phasizes the  effect  of  wavy  radiating  riblets,  as 
described  by  Hanna  and  Smith  for  the  type  lot. 
The  embryonic  sculpture  is  very  similar  to  that 
of  M.  fidelis  (see  Pilsbry  1939). 

The  onset  of  neanic  growth  is  marked  by  an 
abrupt  enlargement  of  whorl  diameter.  The  in- 
tensity of  surface  granulation  on  early  neanic 
whorls  is  less  than  that  on  the  protoconch;  the 
granules  are  sometimes  obscure  or  localized  in 
a  few  patches.  Irregular,  axial  growth  ridges  are 
the  dominant  sculptural  feature.  Between  these 


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FIGURES  6-9.  Figs.  6a-h.  Genitalia  of  Monadenia  troglodytes  wintu,  n.  subsp.,  Loc.  45;  (a.)  dorsal  aspect;  (h.)  ventral 
aspect,  atrium  partially  everted.  Fig.  7.  Copulatory  pad  of  M '.  t.  wintu.  Figs.  &i-b.  Genitalia  of  Monadenia  churchi,  Loc.  29. 
(a.)  dorsal  aspect;  (h.)  detail  of  verge.  Figs.  9a-c.  Ontogenetic  shape  change  in  Monadenia  troglodytes  troglodytes,  Loc.  10. 
Scale  line  =  5  mm  for  Fig.  9<i,  10  mm  for  Figs.  9h,  9c.  Abbreviations  as  in  Figs.  4-5. 


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389 


ridges  there  is  an  extremely  fine,  wrinklelike, 
parallel  lineolation,  particularly  evident  on  the 
base.  Closely  set,  wavy,  spiral  striation  appears 
on  the  base,  usually  by  the  end  of  the  first  post- 
nuclear  whorl.  These  striae  are  irregular  in  both 
strength  and  spacing. 

Variably  expressed,  these  elements — granu- 
lation, growth  rugae,  wrinkle-lineolation,  and 
spiral  striae — constitute  the  microsculpture  of 
adult  shells.  When  not  pitted  or  breached  by 
erosion,  the  periostracum  is  smooth  and  glossy; 
in  adult  shells  it  is  often  scratched  and  abraded 
on  base  and  spire.  Fine,  wavy,  spiral  striation 
on  the  base  is  apparently  standard  but  frequently 
removed  by  abrasion.  On  some  shells,  stronger, 
incised  striae  occur  on  the  shoulder  of  the  last 
whorl  and  may  be  prominent  behind  the  lip. 

After  about  the  three- whorl  stage,  granulation 
is  rare  below  the  shoulder.  It  persists  on  the 
spire  for  a  varying  number  of  whorls,  usually 
becoming  sparser  with  ontogeny,  the  granules 
becoming  lower,  more  elongate,  and  less  clearly 
defined  until,  in  some  cases,  they  finally  merge 
into  the  general  undulation  of  the  periostracal 
surface.  In  most  instances,  however,  granula- 
tion stops  at  or  near  a  growth  rest,  and  when 
growth  is  resumed  the  granules  are  few,  irreg- 
ular, or  absent. 

The  last  granulose  whorl  varies  from  locality 
to  locality  (Fig.  10),  and  there  is  a  half-whorl  to 
two-whorl  range  in  all  samples  large  enough  for 
consideration.  The  means  of  the  last  granulose 
whorl  increase  in  populations  from  north  to 
south  along  the  McCloud  River.  (Far  to  the 
north,  M.  t.  chaceana  has  a  mean  of  1.83  gran- 
ulose whorls,  seemingly  continuing  the  cline.) 
Variation  in  the  Brock  Mountain  herd  is  less 
obviously  clinal:  the  westernmost  samples,  Lo- 
calities 34  and  35,  have  few  granulose  whorls 
(x  =  2.38  and  2.10,  respectively);  the  few  mea- 
surable shells  from  the  more  eastern  localities 
33,  38,  40,  and  41  are  in  the  2.5-3.5  whorl  range; 
and  Locality  39,  situated  geographically  amid 
the  other  eastern  localities,  has  a  predominance 
of  shells  pustulose  at  4.5  whorls  and  beyond 
(x  =  4.17).  Locality  45,  south  of  Shasta  Lake, 
also  has  populations  whose  granulation  persists 
onto  the  later  whorls  (x  =  3.73).  The  difference 
in  number  of  granulose  whorls  between  the 
McCloud  River  and  Brock  Mountain  locality 
groups  is  highly  significant  (P  <  0.001). 

In  hatchlings,  the  protoconch  is  golden  tan; 


MCCLOUD    R. 


IOC    26 
n:  10 

X  =  2  65 


BROCK  MTN. 


I    LOG  33,38,40  41 
n=4 
X=  2.75 
— W — i — ' 


LOG.  45 


CHACEANA 


n-  20 
X  =  3.73 


FIGURE  10.  Histograms  of  last  granulose  whorl  (to  the 
nearest  half  whorl)  in  Monadenia  troglodytes  samples  from 
along  the  McCloud  River  (M.  /.  troglodytes).  Brock  Mountain 
region  and  Loc.  45  south  of  Shasta  Lake  (M.  t.  wintu,  n. 
subsp.),  and  Siskiyou  County  (M.  t.  chaceana). 


some  show  a  faint  reddish-brown  spiral  band  at 
the  periphery  of  the  last  half  whorl.  From  the 
first,  the  teleoconch  shell  material  is  lighter  tan 
than  the  protoconch.  Beginning  with  the  first 
neanic  whorl,  narrow  whitish  zones  border  the 
reddish-brown  band  above  and  below,  the  lower 
zone  seeming  to  appear  earlier  than  the  upper. 
Gradually  the  narrow  band  becomes  darker  and 
more  prominent,  partly  by  contrast  with  the  in- 
creasing whitish  opacity  of  the  rest  of  the  shell. 
The  advancing  suture  partly  or  wholly  obscures 
the  band. 

Color  of  adult  Monadenia  troglodytes  is  vari- 
able, although  fairly  uniform  within  a  sample. 


390 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


The  basic  shell  material  is  white,  covered  in 
fresh,  uneroded  specimens  with  a  transparent, 
yellowish  periostracum.  All  fresh  specimens 
have  a  dark  reddish-brown  band  (chestnut  of 
Rayner  [1970];  Munsell  midpoint  9.5R/2.3/3.8) 
just  above  the  periphery,  ranging  from  0.5  to 
about  1.5  mm  wide  on  the  body  whorl.  Bands 
wider  than  1  mm  occur  mainly  in  the  eastern, 
Brock  Mountain,  samples  and  at  Locality  45 
south  of  Shasta  Lake.  Unpigmented  zones, 
either  white  or  showing  only  the  yellowish  color 
of  the  periostracum,  border  the  dark  suprape- 
ripheral  band  above  and  below.  These  zones  are 
usually  narrower  than  the  dark  band,  but  in 
some  instances  they  are  equally  wide  or  wider. 
The  lower  zone  is  commonly  wider  than  the  up- 
per; on  shells  with  a  light-colored  base,  its  an- 
terior edge  may  be  indistinct. 

Color  of  the  base  is  generally  uniform,  some- 
times slightly  darker  toward  the  periphery.  The 
color  ranges  from  nearly  white  (even  with  peri- 
ostracum intact)  to  a  medium  reddish  brown 
(rust  of  Rayner  [1970];  Munsell  midpoint  9.0R/ 
4.3/8.0),  with  little  variation  within  a  sample. 
The  lightest  bases  occur  in  samples  from  Lo- 
calities 35,  38,  40,  and  41  (all,  Brock  Mountain 
area);  and  the  darkest  at  Localities  39  (Brock 
Mountain)  and  45  (McCloud  limestone  south  of 
Shasta  Lake).  Samples  from  elsewhere  on  the 
McCloud  limestone  are  intermediate  in  shade. 

The  shoulder  and  spire  are  medium  reddish 
brown,  as  dark  as  or  darker  than  the  base.  They 
are  never  as  dark  as  the  supraperipheral  band. 
The  darkest  spires  occur  in  the  Brock  Mountain 
area  and  the  lightest  along  the  McCloud  River. 
In  both  groups  there  is  a  tendency  for  the  center 
of  the  shoulder  to  be  lighter,  producing  a  shad- 
owy secondary  banding  that  is  most  evident  on 
the  lighter  shells  of  the  McCloud  River  herd. 
The  secondary  banding  is  never  as  distinct  as 
that  in  Monadenia  fidelis  (see  Talmadge  1960; 
his  "multibanded  phase").  One  other  color  ef- 
fect is  observable:  some  individuals  have  radial 
streaks  of  darker  pigment  on  the  shoulder. 
These  streaks  usually  precede  a  growth  rest  and 
evidently  represent  concentrated  pigment  de- 
position at  times  when  shell  growth  is  slowing 
down.  They  are  often  followed  by  a  whitish  ra- 
dial streak  (as  growth  starts  up  again  rapidly?). 
When  combined  with  secondary  banding,  as  in 
some  McCloud  River  shells,  the  streaks  produce 
a  mottled  effect. 


It  seems  likely  that  the  same  shell  pigment,  in 
various  dilutions,  produces  the  supraperipheral 
band,  base,  and  shoulder  coloration. 

The  expanded  lip  of  adult  specimens  is  white. 

Color  variation  in  M.  troglodytes  segregates 
geographically:  populations  in  the  Brock  Moun- 
tain area  have  the  darkest  spires,  both  the  dark- 
est and  the  lightest  bases,  and  the  broadest 
bands  near  the  periphery.  Populations  along  the 
McCloud  River  have  the  lightest-colored  shells 
and  the  greatest  incidence  of  secondary  banding 
and  mottling. 

In  the  course  of  ontogeny,  the  shell  of  Mon- 
adenia troglodytes  undergoes  several  significant 
changes  in  shape.  The  protoconch — that  portion 
of  the  shell  which  forms  within  the  egg — is  al- 
most hemispherical,  with  nearly  flat  spire  and 
deeply  convex  base.  Gould  (1969)  suggested  that 
mechanical  limitations  of  space  within  the  egg 
largely  determine  protoconch  shape  in  the  zoni- 
tid  snail  Poecilozonites,  and  in  fact,  the  proto- 
conch of  M.  troglodytes  is  shaped  very  much 
like  that  of  Poecilozonites  bermudensis  bermu- 
densis  (Gould  1969:pl.  6,  figs.  6,  7). 

Post-embryonic  juvenile  shells  are  wide,  flat- 
to  low-spired,  angulate  at  the  shoulder,  and  tu- 
mid at  the  base.  Again  as  in  Poecilozonites, 
postembryonic  growth  begins  with  a  marked  in- 
crease in  relative  width.  As  growth  proceeds, 
the  whorl  changes  in  cross  section  (Fig.  9)  from 
crescentic  and  taller  than  broad,  in  the  embry- 
onic shell,  to  auriculate  and  broader  than  tall  in 
the  adult.  Also  with  growth,  the  periphery, 
which  is  above  the  middle  in  juvenile  shells, 
migrates  downward  until  it  is  medial  in  adults 
and  changes  gradually  from  angulate  to  rounded. 

Spire  height  increases  allometrically  with  re- 
spect to  shell  diameter,  producing  a  domed 
spire.  Since  from  about  the  third  whorl  on,  the 
height-diameter  ratio  of  the  shell  remains  prac- 
tically constant,  the  allometry  is  due  chiefly  to 
the  downward  migration  of  the  periphery. 

In  Monadenia,  as  in  most  other  helicacean 
snails,  growth  is  determinate.  The  shell  enlarges 
up  to  a  point,  generally  coincident  with  sexual 
maturity,  when  the  lip  is  thickened  and  turned 
outward.  Thereafter,  no  additional  spiral  growth 
takes  place.  In  M.  troglodytes,  the  greatest  ab- 
solute height  of  aperture  and  often  the  greatest 
total  shell  height  are  achieved  one-half  to  one- 
quarter  whorl  before  the  cessation  of  growth. 
For  the  last  quarter  turn,  translation  along  the 


ROTH:   DISTRIBUTION.  ANATOMY,  VARIATION  OF  MONADEN1A  TROGLODYTES 


391 


FIGURES  11-13.     Shells  of  Monadenia  troglodytes,  xl.5.  Figs,  lla-c.  M.  t.  troglodytes,  Loc.  26.  Figs,  \2a-c.  M.  t.  wintu, 
n.  subsp.,  holotype  CAS  018431,  Loc.  34.  Figs.  13«-c.  M.  t.  chaceana,  paratype  CASGTC  10125. 


vertical  axis  is  halted  or  reversed,  and  at  the 
same  time,  the  whorl  is  compressed  apico-ba- 
sally.  The  final  3—4  mm  increment  of  growth 
strikes  downward  at  about  a  30°  angle  to  the 
suture,  bringing  the  peristome  nearly  into  tan- 
gency  with  the  face  of  the  body  whorl.  Similar 
terminal  growth  occurs  in  many  genera  of  heli- 
cacean  snails,  particularly  those  from  xeric  en- 
vironments. It  is  presumably  an  adaptation  to 
enhance  the  snails'  mucous  seal  to  the  substra- 
tum, in  order  to  retard  water  loss  or  exclude 
predators.  The  final  half- whorl  "leap"  of  growth 
is  most  strongly  expressed  in  populations  along 
the  McCloud  River  north  of  Shasta  Lake  and 
contributes  to  their  generally  low  height-diam- 
eter ratios  compared  to  Brock  Mountain  popu- 
lations. 

As  already  noted  above.  Monadenia  troglo- 


dytes undergoes  a  much  greater  range  of  variation 
in  shape  than  was  evident  from  the  type  lot 
alone.  Data  on  basic  shape  measurements  and 
indices — height  (H),  diameter  (D),  H/D  ratio, 
number  of  whorls,  umbilical  width  (U).  and  U/D 
ratio — are  summarized  in  Table  3.  Data  from 
adult  shells  (those  with  reflected  lip)  only  are 
included.  Samples  were  collected  without  spe- 
cial procedures  to  insure  randomness,  but  there 
is  no  reason  to  suspect  bias  with  regard  to  any 
of  the  dimensions  or  ratios  used  here. 

All  shells  were  measured  with  a  hand-held  cal- 
iper  with  vernier  scale.  Height  (H)  was  mea- 
sured parallel  to,  and  maximum  diameter  (D) 
perpendicular  to.  the  axis  of  coiling  of  the  shell. 
The  expanded  lip  of  adult  shells  was  excluded 
from  these  dimensions.  Umbilical  width  (U)  is 
an  inside  caliper  measurement  taken  parallel  to 


392 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


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ROTH:   DISTRIBUTION,  ANATOMY.  VARIATION  OF  MONADENIA  TROGLODYTES 


393 


H/D 


.42  .44  .46  .48  .50  .52  .54 


.56  58  .60 


Samwel  types 

Potter  Cr  types 

chaceana 

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28 

43 

45 

35 

31-34,38,40,41 

39 


n:  9 


22 


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U/D 


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10 

11-24 

26,  27 

28 

43 

45 

35 

31-34,38,40,41 

39 


FIGURES  \4a  (upper),  \4b  (lower).  Relative  height  of  spire  (H/D)  and  relative  width  of  umbilicus  (U/D)  in  Monadenia 
troglodytes.  Line  designates  sample  range;  vertical  strike,  mean;  open  bar,  mean  ±  one  standard  deviation;  solid  bar,  95% 
confidence  interval  of  population  mean. 


the  maximum  diameter  and  occasionally  re- 
quired breaking  away  a  portion  of  the  inner  lip. 
Number  of  whorls  was  determined,  with  the  use 
of  a  binocular  microscope,  by  the  method  of 
Diver  (1932)  except  that  the  final  fraction  of  a 
whorl  was  estimated  without  pencil  lines  being 
drawn  on  the  shell.  The  measurement  is  rounded 
to  the  nearest  0.1  whorl.  Pilsbry  (1939:xi,  fig.  B) 
evidently  counted  whorls  by  approximately  the 
same  method,  although  he  usually  expressed  re- 
sults to  the  nearest  one-quarter  or  one-third  of 
a  whorl  and  sometimes  quoted  uncritically  the 
figures  of  other  authors. 

The  simple  measurements  of  size  (H,  D,  H  + 
D)  show  that  as  a  group  the  McCloud  River  herd 
averages  smaller  than  the  Brock  Mountain  herd, 
although  there  is  extensive  overlap  of  ranges  in 
all  categories.  The  smallest  averages  belong  to 
the  samples  from  Localities  10  (below  Samwel 
Cave)  and  28  (Dekkas  Rock).  Shells  of  large  di- 
ameter (and  large  H  +  D)  occur  near  Ellery 
Creek  (Locs.  26,  27). 


Relative  height  of  spire  (H/D)  segregates  geo- 
graphically (Fig.  14«),  with  McCloud  River 
shells  significantly  lower-spired  than  those  from 
Brock  Mountain.  As  in  most  other  parameters, 
shells  from  Locality  45  segregate  with  the  Brock 
Mountain  herd.  The  U/D  ratio,  the  relative 
width  of  the  umbilicus,  segregates  along  the 
same  lines  (Fig.  14/>) — not  surprisingly,  since  the 
two  ratios,  H/D  and  U/D,  may  be  aspects  of  the 
same  variable.  A  tapering  tube  of  a  given  size 
and  rate  of  expansion  may  be  coiled  steeply 
(high  rate  of  translation  along  coiling  axis  and 
small  distance  of  generating  curve  from  axis: 
Raup  1966)  or  flatly  (low  rate  of  translation, 
large  distance  from  axis).  The  first  mode  gen- 
erates a  high-spired  shell  with  small  umbilicus, 
the  second  a  low-spired  shell  with  wide  umbili- 
cus. A  correlation  coefficient  of  -0.951  for  sam- 
ple mean  H/D  and  mean  U/D  further  attests  to 
the  association  between  the  two  parameters  in 
M.  troglodytes  in  the  Shasta  Lake  region.  The 
high  spire  and  small  umbilicus  of  M.  t.  chaceana 


394 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


-  M.t.  chaceana 


31-34, 
38,40,41* 


—  M.t.  wintu 


M.t.  troglodytes  — 


X26,27 


,110  .120 

U/D 


FIGURE  15.  Relation  between  mean  relative  height  of  spire 
(H/D)  and  mean  relative  width  of  umbilicus  (U/D)  in  Mona- 
denia troglodytes. 


41 


39 


37 


35 


33 


31 


'31-34,38.40,41 


26,27^ 


39i 


146 


11-24  * 


'1.4-7 


r43 


5.4 


5.5 


5.6 


5.7 


5.2  5.3 

Whorls 

FIGURE  16.  Relation  between  mean  size  (H  +  D)  and 
mean  number  of  whorls  in  Monadenia  troglodytes.  Diamonds, 
M.  t.  troglodytes;  squares,  M.  t.  wintu,  n.  subsp.;  circle,  M. 
t.  chaceana. 


are  in  harmony  with  this  association,  although 
the  umbilicus  is  broader  and/or  the  spire  higher 
than  regression  of  the  Shasta  Lake  statistics  pre- 
dicts (Fig.  15). 

Figure  16  plots  the  relationship  between  mean 
size  (H  +  D)  and  whorl  number.  Although  the 
scatter  of  individual  (specimen)  points  overlaps 
extensively,  again  there  is  geographic  segrega- 
tion of  means.  Locality  45  sorts  with  the  Brock 
Mountain  herd,  nearest  to  the  pustulose  shells 
from  Locality  39.  Mean  size  and  mean  whorl 
number  correlate  strongly  in  the  McCloud  River 
herd  (r  =  0.954);  the  more  heterogeneous  Brock 
Mountain  herd  displays  no  such  tight  correla- 
tion. 

Based  on  the  dimensions  given  by  Hanna  and 
Smith  (1933),  the  type  lot  of  M.  troglodytes  from 
Samwel  Cave  and  their  material  from  Potter 
Creek  Cave  are  flatter  (lower  H/D)  than  any  of 
the  samples  here  studied.  Both  the  mean  diam- 
eter and  mean  H/D  ratio  of  the  type  lot  differ 
significantly  from  those  of  the  nearest  living 
sample,  from  Locality  10.  It  would  be  prema- 
ture, however,  to  conclude  that  the  difference 
represents  evolutionary  change.  Hanna  and 
Smith  accepted  a  Pleistocene  age  for  the  cave 
specimens,  first  because  they  believed  the 
species  extinct  and,  second,  because  the  shells 
had  been  associated  with  bone  deposits.  (The 
Samwel  Cave  and  Potter  Creek  Cave  vertebrate 


faunas  have  been  assigned  a  Rancholabrean  [late 
Pleistocene]  age;  Hibbard  et  al.  1965.)  But  the 
burrowing  or  crevice-seeking  habit  of  land  snails 
makes  them  unreliable  stratigraphically.  State  of 
preservation  is  not  very  informative:  in  the  sev- 
eral cave  samples  now  at  hand,  there  are  shells 
in  every  gradation  from  slightly  dusty  to  heavily 
encrusted  with  calcium  carbonate,  and  there 
may  be  a  more  or  less  continual  influx  of  shells 
drifting  or  washing  into  the  caves.  Independent 
evidence  of  age,  such  as  radiometric  dating,  and 
collections  with  good  stratigraphic  control  are 
needed  before  one  can  place  the  cave  samples 
in  a  time  framework. 

SYSTEMATICS 

TAXONOMIC  CRITERIA  IN  Monadenia. — In- 
dividuals of  Monadenia  troglodytes  in  the  Shas- 
ta Lake  region  group  strongly  into  two  geo- 
graphic units  based  on  shell  color  and  texture, 
microsculpture,  and  morphometry.  Monadenia 
chaceana  constitutes  a  third  such  unit.  Although 
its  internal  anatomy  is  not  yet  known,  its  shell 
characters  are  highly  similar  to  M.  troglodytes, 
and  it  is  thus  given  coordinate  rank  with  the  lat- 
ter two  groups. 

The  subspecies  category  has  been  employed 
somewhat  unevenly  in  terrestrial  malacology. 
Early  workers  responded  to  the  manifold  vari- 
ability of  some  tropical  snail  groups,  for  in- 


ROTH:  DISTRIBUTION,  ANATOMY,  VARIATION  OF  MONADENIA  TROGLODYTES 


395 


stance,  by  naming  dozens  of  subspecies,  and  the 
morass  of  names  led  in  turn  to  a  reaction  against 
taxonomic  recognition  of  infraspecific  units 
(Gould  1969).  I  agree  with  Kavanaugh  (1979:93) 
that  "recognition  of  the  usefulness  of  a  subspe- 
cies concept  .  .  .  [depends]  on  one's  particular 
perspective;  more  specifically,  on  the  distribu- 
tion of  habitats  or  areas  occupied  by  the  organ- 
isms one  studies.  Where  gaps  between  areas  of 
suitable  habitat  are  broad  and  clear,  .  .  .  corre- 
lated discontinuity  in  variation  is  more  confi- 
dently recognized  and  suggestive  of  active,  ef- 
fective barriers  to  gene  flow."  The  three  groups 
of  Monadenia  troglodytes  meet  the  criteria  for 
subspecies.  Shell  character  differences  are 
mainly  quantitative  and  correlated  with  geog- 
raphy. Except  for  the  different  verge  configu- 
ration at  Locality  43  (the  significance  of  which 
remains  unresolved),  the  identity  of  the  genitalia 
between  the  two  Shasta  Lake  region  herds  sug- 
gests that  interbreeding  would  be  possible  if  the 
two  were  brought  together  sympatrically  in  na- 
ture. Limestone  substratum  is  apparently  the 
required  habitat,  its  absence  a  probable  barrier 
to  gene  flow,  at  least  under  present  environmen- 
tal conditions.  Rivers  are  at  least  short-term  bar- 
riers, but  over  the  long  term  may  be  important 
agents  of  dispersal.  The  population  at  Locality 
45  may  well  have  been  established  by  river- 
borne  waifs  rafted  down  the  Pit  River — perhaps 
from  the  population  of  Locality  39,  with  which 
it  groups  morphologically  (Figs.  10,  14-16).  Cli- 
nal  variation  in  granulation  along  the  McCloud 
River  arm  suggests  incomplete  genetic  isolation 
between  those  localities.  Populations  on  the 
McCloud  limestone,  exclusive  of  Locality  45, 
constitute  one  unit;  since  it  includes  the  type- 
locality,  this  unit  is  the  nominate  subspecies, 
Monadenia  (Shastelix)  troglodytes  troglodytes. 
Populations  between  the  Pit  River  and  Squaw 
Creek  arms  of  Shasta  Lake,  and  the  southern 
population  at  Locality  45,  are  named  as  a  new 
subspecies  below. 

The  sympatric  or  parapatric  occurrence  of 
Monadenia  churchi  and  M.  troglodytes  at  sev- 
eral localities  in  the  Shasta  Lake  region  should, 
theoretically,  permit  an  estimate  of  the  degree 
of  morphologic  difference  that  can  be  expected 
between  species  of  the  genus,  and  by  analogy 
aid  in  ranking  allopatric  taxa.  Reproductive 
structures  are  particularly  suitable  for  such  an 
analysis,  because  a  major  difference  in  genitalia 


(taking  into  account  the  possible  effects  of  sea- 
sonal variation  and  genital  polymorphism)  im- 
plies functional  incompatibility.  Solem  (1975) 
used  data  on  reproductive  anatomy  of  sympatric 
species  to  define  taxonomic  criteria  in  the  snail 
genus  Oreohelix  Pilsbry,  1904.  Like  many  other 
helicacean  snails,  monadenias  have  elaborate 
terminal  genitalia,  replete  with  accessory  organs 
which  are  employed  in  a  courtship  of  some  com- 
plexity (Webb  1952,  1966).  Specialized  repro- 
ductive organs  and  complex  precopulatory  be- 
havior are  sometimes  regarded  as  isolating 
mechanisms  in  other  animal  groups;  theoreti- 
cally, they  prevent  unproductive  mating  be- 
tween sympatric  species.  Solem  (1978:67)  pre- 
dicted that,  among  pulmonates,  elaboration  of 
stimulatory  and  glandular  or  dart  structures 
would  occur  where  the  need  for  species  recog- 
nition signs  was  the  greatest,  for  example,  under 
conditions  of  sympatry  in  areas  of  historically 
fluctuating  climate  and  vegetational  cover  or  in 
island  situations  involving  explosive  speciation. 
The  differences  in  the  genitalia  of  M.  churchi 
(Fig.  8)  and  M.  troglodytes  (Figs.  4-6),  how- 
ever, are  relatively  limited  and  far  less  marked 
than  those  between  members  of  different  sub- 
genera. 

The  values  for  organ  lengths  in  M.  churchi 
(Table  2)  fall  mostly  within  the  range  of  variation 
of  M.  troglodytes.  The  dart  sac  averages  smaller 
in  M.  churchi,  but  the  size  range  overlaps  that 
of  M.  troglodytes.  Length  of  the  mucus  gland 
is  highly  variable  in  both  species  and  may  vary 
according  to  recency  of  copulation.  M.  churchi 
from  Mammoth  Butte,  west  of  Shasta  Lake,  and 
others  from  the  Trinity  River  drainage  have  a 
thin,  cylindric  verge  with  the  end  squared  and 
compressed,  the  meatus  transverse.  The  verge 
of  a  specimen  from  near  Campbell  Creek  on  the 
east  side  of  the  McCloud  River  arm  (Loc.  29)  is 
also  squared  at  the  tip  but  thicker,  with  lateral 
meatus  and  concave  facet  as  in  M.  troglodytes 
(Fig.  86).  All  specimens  examined  have  a  single 
large  pilaster  in  the  penial  chamber.  It  appears, 
therefore,  that  reproductive  anatomy  in  Shas- 
telix  is  too  variable  and  too  weakly  differentiated 
to  offer  reliable  characters  for  diagnosis. 

The  two  species  are  much  more  readily  dis- 
tinguished on  shell  characters:  M.  churchi  has 
strong,  pustulose  microsculpture  over  the  entire 
shell,  even  when  adult;  a  dull  periostracal  sur- 
face; and  uniform  brown  color  on  base  and 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


spire.  Quantitative  differences,  not  always  pres- 
ent but  usually  helpful  in  distinguishing  the  two 
species,  include  higher  spire,  stronger  growth 
rugae,  and  more  tightly  coiled  last  whorl. 

I  have  detected  no  systematic  conchological 
differences  between  M.  troglodytes  found  with 
M.  churchi  and  those  occurring  alone.  In  each 
case  the  M.  troglodytes  shells  retain  the  stan- 
dard characters  of  their  herd.  On  the  other  hand, 
M.  churchi  from  within  the  range  of  M.  trog- 
lodytes tend  to  be  larger,  shinier,  more  inflated, 
and  less  densely  pustulose  than  specimens  from 
elsewhere.  Whether  this  apparent  convergence 
with  M.  troglodytes  goes  beyond  independent 
geographic  variation  will  be  examined  in  a  later 
paper. 

The  finding  that  in  reproductive  anatomy  M. 
churchi  and  M.  troglodytes  differ  from  each  oth- 
er chiefly  in  verge  shape  also  implies  that  the 
elaborations  of  the  female  side  of  the  genital  sys- 
tem— dart  sac,  copulatory  pad,  etc. — may  not 
be  involved  in  species  recognition  during  court- 
ship or  maintenance  of  reproductive  isolation. 
According  to  the  sexual  selection  model  of 
Charnov  (1979),  hermaphroditic  animals  may 
copulate  not  so  much  to  gain  sperm  to  fertilize 
eggs  as  to  give  sperm  away  (to  gain  access  to 
another's  eggs).  "There  must  often  exist  a  con- 
flict of  interest  between  mating  partners — as  a 
recipient  each  should  be  inclined  to  accept 
sperm  (not  necessarily  for  fertilization  of  its  own 
eggs)  in  order  to  give  its  sperm  away"  (Charnov 
1979:2482).  In  order  to  pair  with  and  fertilize  a 
partner,  a  Monadenia  may  have  to  display  its 
own  apparent  receptivity  to  the  partner's 
sperm.  Like  any  character  that  individuals  use 
to  choose  sperm  donors,  the  organs  involved  in 
such  a  display  would  be  subject  to  exaggeration 
through  time.  Selective  pressure  toward  species- 
specific  differentiation  of  the  organs  would  de- 
pend on  (1)  frequency  of  the  opportunity  for  in- 
terspecific mating,  based  both  on  degree  of  sym- 
patry  and  the  effectiveness  of  other  organ  or 
behavior  systems  in  maintaining  isolation;  and 
(2)  relative  fitness  of  resulting  hybrids. 

Monadenia  (Shastelix)  troglodytes  troglodytes 

Hanna  and  Smith 

(Figures  4<i-b,  9a-c,  ll«-c) 

Monadenia  troglodytes  HANNA  AND  SMITH,  1933:84-85,  pi. 
5,  figs.  6-8.— PIISBRY  1939:54,  fig.  22(6-8).— SMITH 
1957:26;  1970:40.— WAI  TON  1970: 1 1 1.— ROTH  1972a:7: 
1972b:7;  1979:13. 


ORIGINAL  DESCRIPTION. — "Shell  light  buff, 
medium  size,  widely  umbilicate;  spire  greatly 
depressed;  whorls  5Vi  with  moderately  deep  su- 
ture; the  last  whorl  slightly  depressed  near  the 
aperture;  outer  margin  expanded  very  little,  the 
basal  margin  somewhat  more  so;  one  pale  brown 
spiral  band  appears  just  above  the  periphery, 
which  is  bounded  above  and  below  by  white 
bands  that  are  slightly  wider;  surface  without 
markings  except  growth  lines;  nucleus  consist- 
ing of  \Vi  whorls  marked  by  radiating  wavy  rib- 
lets.  Diameter  24.2;  altitude  10.8  mm"  (Hanna 
and  Smith  1933).  This  description  was  repeated 
by  Pilsbry  (1939). 

TYPE  MATERIAL.— Holotype:  UCMP  32394  (shell):  CALI- 
FORNIA: Shasta  County:  Samwel  Cave  (UCMP  loc.  1008). 

Paratype:  CASGTC  5842  (shell);  same  locality  as  holotype. 

REFERRED  MATERIAL. — CALIFORNIA:  Shasta  County:  Bol- 
libokka  Mountain  Quadrangle:  Loc.  8  [2  specimens],  9  [5],  10 
[32],  11  [6  in  addition  to  holotype  and  paratype],  12  [2],  13  [2], 
14  [2],  15  [2],  16  [1],  17  [4],  18  [2],  19  [31],  20  [2],  21  [6],  22 
[1],  23  [2],  24  [9],  25  [2],  26  [30],  27  [5],  28  [38],  29  [3].  Lamoine 
Quadrangle:  Loc.  42  [1],  43  [22],  44  [12],  47  [1],  48  [1].  Figure 
3  maps  these  localities. 

DISCUSSION. — As  described  above  in  the  sec- 
tion on  geographic  variation  and  graphically 
shown  in  Figures  14-16,  M.  t.  troglodytes  has 
the  lowest-spired  and  most  broadly  umbilicate 
shells  of  any  subspecies.  The  shells  average 
smaller  than  those  of  the  Brock  Mountain  herd 
(M.  t.  wintu,  next  described),  but  large  shells 
occur  near  Ellery  Creek.  The  lightest-colored 
shells  and  the  greatest  incidence  of  secondary 
banding  and  mottling  occur  in  M.  t.  troglodytes. 
The  nominate  subspecies  tends  to  have  fewer 
coils  to  the  epiphallus  than  M.  t.  wintu. 

Monadenia  (Shastelix)  troglodytes  wintu,   new 

subspecies 

(Figures  5a-b,  dti-b,  7,  \2a-c) 

DIAGNOSIS. — M.  troglodytes  with  moderately 
high  spire  (sample  mean  H/D  =  0.526-0.546), 
shell  solid,  lustrous,  shoulder  reddish  brown, 
strong  white  zones  above  and  below  brown  su- 
praperipheral  band,  base  either  nearly  white,  or 
reddish  brown,  as  dark  as  shoulder.  Granulation 
often  persisting  past  third  whorl. 

DESCRIPTION. — Shell  of  moderate  size  for  the 
genus,  solid,  moderately  to  distinctly  thick,  lus- 
trous; spire  convexly  conic,  moderately  elevat- 
ed; whorls  5.2-6.1  in  adult,  tightly  coiled;  body 
whorl  slightly  expanded  over  last  Vz  turn.  Pro- 
toconch  of  1.5-1.9  whorls,  nuclear  tip  smooth, 


ROTH:  DISTRIBUTION.  ANATOMY.  VARIATION  OF  MONADEN1A  TROGLODYTES 


397 


followed  by  fine  granulation  tending  to  form 
wavy,  radial  riblets  below  suture;  granules  else- 
where spirally  elongated,  in  diagonal  series  or 
irregularly  scattered.  Early  neanic  whorls  less 
densely  granulated,  granulation  becoming 
sparser  with  growth,  usually  persisting  past  the 
2. 5- whorl  stage  and  often  past  the  3- whorl  stage 
in  at  least  some  members  of  each  sample.  Body 
whorl  sculptured  with  fine  growth  rugae  and 
microscopic,  wavy,  parallel,  spiral  lineolation, 
most  evident  behind  lip.  Juvenile  shells  with 
base  tumid,  spire  low,  shoulder  angulate;  pe- 
riphery becoming  first  obtusely  angular  and  fi- 
nally rounded  with  maturity.  Base  of  adult  shells 
rounded,  umbilicus  open,  steep- walled,  diame- 
ter 0.090-0.155  times  major  diameter  of  shell. 
Last  3-4  mm  of  body  whorl  striking  downward 
at  about  30°  angle  to  suture.  Aperture  broadly 
auriculate,  apico-basally  compressed,  oblique. 
Peristome  slightly  thickened,  everted;  inner  lip 
covering  10-25%  of  umbilicus.  Color  of  spire 
and  shoulder  medium  reddish  brown  (rust  of 
Rayner  [1970];  Munsell  midpoint  9.0R/4. 3/8.0); 
dark  reddish  brown  (chestnut  of  Rayner  [1970], 
Munsell  midpoint  9.5R/2.3/3.8)  band  just  above 
periphery,  1-1.5  mm  wide  in  adult,  bordered 
above  and  below  with  white  or  light  tan  zones, 
upper  zone  either  narrower  or  about  as  wide  as 
brown  band,  lower  zone  usually  as  wide  or 
somewhat  wider;  base  whitish  with  light  tan  suf- 
fusion or  (at  some  localities)  reddish  brown  as 
dark  as  shoulder.  Periostracum  smooth,  color- 
less or  very  light  yellowish  tan.  Dimensions:  ho- 
lotype,  height  12.8  mm,  diameter  25.1  mm,  di- 
ameter of  umbilicus  3.0  mm,  5.6  whorls;  largest 
paratype  (Loc.  35),  height  14.9  mm,  diameter 
27.6  mm,  diameter  of  umbilicus  2.9  mm,  5.6 
whorls.  Dimensions  of  other  referred  material 
summarized  in  Table  3. 

Body  of  animal  dove  gray  to  sooty  black  with 
a  reddish  or  purplish  cast,  darker  on  dorsum; 
light  mid-dorsal  stripe;  sole  light  buff  with  gray 
margin.  Mantle  over  lung  translucent  buff  with 
gray  pencilling  covering  less  than  10%  of  sur- 
face. Genitalia  (Figs.  5,  6)  as  in  typical  subspe- 
cies. 

Jaw  as  in  M.  t.  troglodytes.  Radula  substan- 
tially as  in  M.  t.  troglodytes,  with  42  teeth  in  a 
half  row  (Locs.  34,  45),  an  endocone  developed 
on  tooth  21  (Loc.  34)  or  14  (Loc.  45),  an  ecto- 
cone  on  tooth  22  (Loc.  34)  or  17  (Loc.  45).  Bifid 
and  trifid  endocones  occur  sporadically. 


TYPE  MATERIAL.— Holotype:  CAS  018431  (shell,  radula, 
and  soft  parts);  CALIFORNIA:  Shasta  County:  NW'/i  NE14  sec. 
8,  T.  34  N,  R.  2  W,  USGS  Bollibokka  Mountain  Quadrangle 
(15-minute  Series  [Topographic];  ed.  1957),  cave  between  two 
limestone  buttes  at  south  end  of  Gray  Rocks,  above  Pit  River 
arm  of  Shasta  Lake.  S.  E.  Hirschfeld  coll.,  Oct.  1975.  [Loc. 
34.] 

Paratypes:  CAS  018432,  018433  (shells  and  soft  parts),  same 
locality  as  holotype.  CAS  018434,  16  shells,  same  locality  as 
holotype.  CAS  018435,  5  shells,  Loc.  35.  USNM,  ANSP, 
AMNH,  FMNH,  LACM,  one  shell  each,  Loc.  35. 

REFERRED  MATERIAL. — CALIFORNIA:  Shasta  County:  Bol- 
libokka Mountain  Quadrangle:  Loc.  30  [2  specimens],  31  [19], 
32  [8],  33  [1],  36  [3],  37  [5],  38  [7],  39  [13],  40  [14],  41  [20]. 
Project  City  Quadrangle:  Loc.  45  [40].  Figure  3  maps  these 
localities. 

ETYMOLOGY. — The  subspecies  is  named  for 
the  people  native  to  the  region  where  it  is  found, 
the  Wintu  tribe. 

DISCUSSION. — The  major  features  of  variation 
within  the  subspecies  and  its  morphological  re- 
lationship to  other  subspecies  are  discussed 
above  and  graphically  shown  in  Figures  14-16. 
In  coloration  and  spire  height,  M.  t.  wintu  and 
M.  t.  chaceana  are  more  similar  than  either  is 
to  M.  t.  troglodytes,  whereas  in  microsculpture 
and  relation  of  whorl  number  to  overall  size,  M. 
t.  chaceana  and  M.  t.  troglodytes  group  more 
closely. 

Monadenia  (Shastelix)  troglodytes 
chaceana  Berry 

(Figures  13«-c) 

Monadenia  chaceana BERRY,  1940b:9-ll,  figs  9,  10. — PILSBRY 
1948:1092.— SMITH  1960:97.— ROTH  1972a:5;  1972b:6. 

ORIGINAL  DESCRIPTION. — "Shell  of  but  mod- 
erate size,  weight,  and  thickness;  spire  low-con- 
ic to  moderately  elevated;  whorls  5Vz  to  6, 
subangulate  and  carinate  above  the  middle 
during  juvenility,  subcarinate  at  adolescence, 
but  becoming  obtusely  angular  and  finally  quite 
well  rounded  at  maturity;  base  tumid,  the  um- 
bilicus open,  steep-walled,  permeable  to  apex, 
and  contained  on  the  average  about  8.4  times 
(7.45  to  9.88  in  those  measured)  in  the  major 
shell-diameter.  Aperture  somewhat  descending 
above,  oblique,  rounded  to  round-ovate,  slightly 
or  not  at  all  flattened  below;  peristome  nearly 
simple  above,  elsewhere  usually  little  thickened 
and  but  moderately  everted,  terminating  below 
in  a  very  moderate  columellar  flare  which  covers 
only  the  edge  of  the  umbilicus. 

"Embryonic  shell  swollen,  of  1%  to  2  whorls; 
the  surface  initially  smooth,  but  almost  at  once 
breaking  into  a  few,  irregular  axial  waves  sue- 


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ceeded  by  a  close,  fine,  crowded  granulation, 
which  abruptly  ceases  with  the  first  post-embry- 
onic whorl,  the  often  heavy  but  extremely  vari- 
able growth-striae  thenceforth  becoming  the 
dominant  feature;  a  few  indistinct,  elongate,  and 
commonly  confluent  papillae  arranged  in  rather 
distant  forward-descending  series  appear  on  the 
early  turns,  but  gradually  give  way  on  the  later 
whorls  to  a  weak  and  not  very  regular  spiral 
striation  which  may  become  quite  indistinct  on 
the  base;  general  surface  between  the  striae  and 
growth-lines  very  finely  microscopically  wrin- 
kled in  a  cloth-like  pattern. 

"Periostracum  smooth  and  lustrous;  deep 
brown,  encircled  by  a  conspicuous  dark-brown 
band  about  2  mm.  wide  just  above  the  periphery, 
bordered  by  a  much  narrower  yellowish  band 
and  yet  narrower  band  of  the  same  pale  tone  just 
above,  while  on  the  shoulder  some  shells  show 
varying  traces  of  yet  another  band  of  interme- 
diate brownish  tone,  best  seen  in  juvenals" 
(Berry  1940b). 

TYPE  MATERIAL.— Holotype:  SSB  8678  (shell);  CALIFOR- 
NIA: Siskiyou  County:  among  rocks  about  halfway  up  a  spur 
of  Badger  Mountain  on  west  side  of  Shasta  River  Canyon  not 
far  above  its  mouth  (Berry  1940b). 

Paratypes:  CASGTC  10125  (formerly  A.  G.  Smith  no.  7102; 
shell);  SUPTC  6555  (shell);  according  to  Berry  (1940b),  ad- 
ditional paratypes  are  in  the  Berry  Collection,  with  others 
originally  to  be  deposited  in  the  USNM  and  the  Emery  P. 
Chace  Collection. 

DISTRIBUTION. — Monadenia  t.  chaceana  is 
known  only  from  the  general  vicinity  of  the  type- 
locality,  in  the  Hornbrook  Quadrangle,  Siskiyou 
County  (Locs.  1-5),  and  from  one  site  (Loc.  6) 
in  the  Yreka  Quadrangle,  Siskiyou  County  (see 
also  Fig.  1).  Locality  7,  "near  Yreka"  is  too 
generalized  to  assign  to  a  quadrangle  with  con- 
fidence. Limestone  areas  west  of  Gazelle  and 
southeast  of  Scott  Valley  need  additional  pros- 
pecting for  Monadenia. 

DISCUSSION. — In  coloration,  M.  t.  chaceana 
most  closely  resembles  the  darker  samples  of 
M.  t.  wintu.  The  base  is  as  dark  as  the  spire, 
although  neither  is  as  dark  as  the  supraperiph- 
eral  band.  At  Localities  5  and  7,  along  with 
the  usual  dark-spired  individuals,  a  form  is  pres- 
ent with  light  shoulder  and  one  narrow,  medium- 
brown,  secondary  band  about  two-thirds  the  dis- 
tance from  the  suture  to  the  periphery. 

The  substance  of  the  shell  is  thinner  and  more 
translucent  than  in  the  Shasta  Lake  region 
groups. 


Monadenia  troglodytes  chaceana  has  the 
highest  mean  H/D  ratio  of  any  subspecies  (Table 
3,  Fig.  \4a),  differing  highly  significantly  from 
any  sample  of  M.  t.  troglodytes  or  M.  t.  wintu. 
At  equivalent  diameters,  adult  M.  t.  chaceana 
average  0.49  whorl  more  than  M.  t.  wintu  from 
the  Brock  Mountain  area. 

Berry  (1940b)  compared  his  M.  chaceana  to 
Monadenia  mormonum  cala  (Pilsbry,  1900)  and 
M.  churchi.  He  also  intimated  a  possible  rela- 
tionship to  Monadenia  fidelis  minor  (Binney, 
1885)  and  surmised  that  the  type-locality  of  that 
subspecies  might  be  the  Mount  Shasta  region. 
The  latter  supposition  was  evidently  based  on 
a  statement  by  Binney  (1885;  Binney  and  Bland 
1869)  that  small  Monadenia  fidelis  occur  at 
Mount  Shasta. 

Binney  (1885:121,  fig.  91)  proposed  "Aglaia 
fidelis  var.  minor"  without  a  diagnosis  and  his 
original  figure  is  unlocalized,  but  elsewhere 
(1885:141,  footnote;  493)  he  recorded  it  from 
The  Dalles,  Oregon,  collected  by  Henry  Hemp- 
hill.  Henderson  (1936)  and  Pilsbry  (1939)  ac- 
cepted The  Dalles  as  the  type-locality.  The  Cal- 
ifornia Academy  of  Sciences  collection  contains 
two  probable  syntypes  of  A.  f.  var.  minor, 
CASGTC  6001-6002,  formerly  H.  Hemphill  Col- 
lection no.  8598  and  labeled  "Helix  fidelis  Gray,/ 
var.  minor  W.  G.  Binn/depressed  near  mormon- 
um/Near  The  Dalles,  Oregon/Types/HH"  in 
Hemphill's  handwriting.  They  are  small  M.  fi- 
delis, agreeing  well  with  Pilsbry's  (1939)  diag- 
nosis of  M.  fidelis  minor  and  moderately  well 
with  Binney's  original  engraving.  That  figure 
shows  a  pale  shoulder  with  a  light-centered  spi- 
ral band  and  an  indication  of  strong  radial 
growth  striae  on  the  early  whorls — all  charac- 
teristic of  the  Dalles  form. 

The  Binney  and  Bland  Collection  of  terrestrial 
mollusks,  now  at  the  American  Museum  of  Nat- 
ural History  (Gratacap  1901),  contains  one  spec- 
imen of  M.  fide  Us  labeled  "H.  fidelis/Mt.  Shasta/ 
Cooper."  An  oval,  gold-edged,  adhesive  label 
on  the  specimen  states  "Mt.  Shasta/WHB 
JGC";  the  shell  was  evidently  collected  by  Wil- 
liam H.  Brewer  and  given  to  James  G.  Cooper, 
who  passed  it  on  to  Binney.  This  specimen, 
AMNH  57788,  is  25.8  mm  in  diameter,  16.1  mm 
high,  with  6.0  whorls.  The  shoulder  is  unpig- 
mented  except  for  faint,  discontinuous  traces  of 
a  light-centered  spiral  band.  It  is  not  the  speci- 
men illustrated  by  Binney  (1885). 


ROTH:  DISTRIBUTION.  ANATOMY,  VARIATION  OF  MONADENIA  TROGLODYTES 


399 


To  my  knowledge,  M.fidelis  does  not  live  at 
Mt.  Shasta  proper,  the  nearest  localities  being 
Beaver  Creek  near  the  Klamath  River,  Siskiyou 
County  (subspecies  M.f.  leonina  Berry,  1937), 
and  near  the  shores  of  Klamath  Lake,  Oregon 
(unnamed  subspecies  cited  by  Pilsbry  [1939:42, 
fig.  18e]  as  M.f.  minor}.  The  Binney  specimen 
is  similar  to  the  latter  but  more  narrowly  umbil- 
icate.  Brewer  probably  collected  this  shell  on 
his  1863  trip  through  the  Klamath  Mountains, 
rather  than  on  his  1862  climb  of  Mount  Shasta 
(Brewer  1930).  Cooper  (1869)  mentioned  receiv- 
ing from  Brewer  M.fidelis  collected  at  Crescent 
City,  the  western  terminus  of  Brewer's  Klamath 
route.  I  conclude  (1)  that  the  name  minor  applies 
to  a  race  of  small  Monadenia  fidelis  (Gray)  from 
around  The  Dalles,  Oregon,  and  (2)  that  Berry 
was  mistaken  about  the  similarity  of  his  M.  cha- 
ceana  to  Binney's  "Mount  Shasta"  material. 

PHYLOGENETIC  HYPOTHESES 
CONCERNING  Monadenia 

Early  statements  about  evolution  within  Mon- 
adenia tended  to  accord  the  widespread  and 
conspicuous  M.  fidelis  a  central — and  in  some 
unspecified  way  archetypal — position,  while  the 
smaller,  more  remote  and  cryptic  M.  hille- 
brandi,  M.  mormonum,  and  similar  forms  were 
viewed  as  derivative  (and  possibly  degenerate) 
offshoots  (for  example,  see  Cooper  1887; 
Stearns  1900).  A  questionable  exception  is  that 
of  Cooper  (1873),  who  evidently  attempted  to 
adduce  general  principles  for  the  direction  of 
character-state  transformation  in  west  coast 
snail  genera.  One  can  readily  speculate  as  to 
what  extent  an  ethnocentric  outlook  colored 
such  views.  A  second  generation  of  malacolo- 
gists  studying  Monadenia  (chief  among  them 
Pilsbry,  S.  S.  Berry,  G  D.  Hanna,  and  Junius 
Henderson)  concerned  themselves  more  with 
description  and  less  with  interpretation  of  evo- 
lutionary relationships.  But  in  their  work  too,  a 
typological  bias,  with  M.  fidelis  the  "type"  in 
more  than  a  nomenclatural  sense,  may  be  de- 
tected. In  a  recently  proposed  alternative  hy- 
pothesis (Roth  1979),  the  ancestral  Monadenia 
was  seen  as  a  ground-dwelling,  low-spired,  and 
somewhat  variable  form,  from  which  the  ex- 
posed-crawling, partly  diurnal,  and  semi-arbo- 
real M.  fidelis  evolved:  "Once  emancipated 
from  life  in  holes  in  the  ground,  the  fidelis  group 
achieved  large  size,  relatively  high  spire,  and  an 


TABLE  4.     CHARACTER  COMPLEXES  IN  Monadenia  (apo- 
morphous  states  listed  first). 

1.  Atrium  small  and  narrow  (large  and  globose). 

2.  Mucus  gland  about  equal  to  (many  times  longer  than)  dart 
sac. 

3.  Lower  part  of  mucus  gland  adnate  to  (free  from)  atrium. 

4.  Dart  sac  5  mm  or  less  in  length  (longer  than  5  mm). 

5.  Flagellum  about  as  long  as  (^1.5  times  length  of)  penis 
plus  epiphallus. 

6.  Flagellum  and  spermatophore  straight  (helically  coiled). 

7.  Basal  chamber  of  penis  sessile  on  (invaginated  into) 
atrium. 

8.  Penial  retractor  inserted  near  middle  (on  distal  third)  of 
epiphallus. 

9.  Habit  at  least  partly  arboreal  (ground-dwelling). 

10.  Granulose  microsculpture  present  on  teleoconch  (limited 
to  protoconch). 

11.  Shoulder  band   pigmentation   monomorphic  (polymor- 
phic). 


extensive  range  which  now  reaches  farther  north 
than  any  other  American  helicacean"  (Roth 
1979:13).  Roth  and  Eng  (1980)  offered  a  conjec- 
tural, "narrative"  (sensu  Ball  1976)  hypothesis 
for  the  origin  of  Monadenia  setosa  Talmadge. 
Such  narrative  explanations  are  not  analytical 
and  have  little  predictive  power.  To  date,  none 
of  these  competing  models  has  specified  the  cri- 
teria for  judging  relationships,  and  none  has 
been  couched  in  testable  propositions. 

Phylogenetic  systematics,  the  methodology  of 
Willi  Hennig  (1966),  seeks  to  analyze  the  prob- 
able direction  of  evolutionary  character  trans- 
formation in  a  particular  group  of  organisms 
and,  from  this  analysis,  to  reconstruct  the  evo- 
lutionary history  of  the  group.  An  important  vir- 
tue of  this  method  is  that  it  generates  testable 
hypotheses  about  phylogenetic  relations.  (See 
Kavanaugh  [1972]  for  an  exposition  of  Hennig's 
approach  to  systematics  and  Gaffney  [1979]  for 
a  useful  bibliography  of  papers  dealing  with  cla- 
distic  methodology.)  In  land  malacology,  Van 
Goethem  (1977),  Breure  (1979),  and  Bishop 
(1979)  have  applied  these  principles  to  the  land 
snail  groups  Urocyclinae,  Bulimulidae,  and 
American  Camaenidae,  respectively,  and  the 
same  approach  is  implicit  in  the  writings  of  a 
number  of  other  malacologists. 

Of  55  morphological  and  natural  history  char- 
acters studied,  11  (Table  4)  were  incorporated 
in  a  cladistic  analysis.  Characters  excluded  were 
those  in  which  polarity  of  the  transformations 
could  not  be  interpreted,  those  that  occur  only 


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Monadenia,  s.  s  Shastelix  Corynadenia 


5,6,9 


1.2.11 


FIGURE  17.  Suggested  cladogram  for  the  subgenera  of 
Monadenia.  Numbers  refer  to  apomorphous  states  of  char- 
acter complexes  listed  in  Table  4. 


in  states  judged  to  be  symplesiotypic  for  the  ge- 
nus, and  those  that  are  unique  (as  apomorphies) 
to  one  species  and  hence  provide  no  information 
for  grouping. 

Figure  17  illustrates  the  relationship  of  char- 
acter states  in  the  three  subgenera  of  Monaden- 
ia. It  is  well  to  emphasize  that  the  hypothesis 
presented  here  is  preliminary  and  undoubtedly 
will  be  modified  as  study  of  the  genus  pro- 
gresses. This  cladogram  is  based  mainly  on  re- 
productive anatomy.  In  general,  shell  characters 
are  much  more  labile,  phenotypic  responses  to 
environment  are  common,  and  the  number  of 
conchologically  polytypic  species  in  Monadenia 
hampers  the  use  of  shell  features  in  this  kind  of 
analysis.  As  Bishop  (1979:275)  noted,  snail 
shells  offer  relatively  few  characters  for  analysis 
and  are  difficult  to  characterize  objectively, 
"though  both  shape  and  texture  may  be  most 
informative  to  the  eye  of  an  experienced  worker 
in  a  manner  which  is  hard  to  express  in  words." 
Moreover,  shell  characters  are  subject  to  nu- 
merous parallelisms  and  convergences,  usually 
associated  with  habit  and  habitat  (for  example, 
the  often-remarked  similarity  between  the  rock 
crevice-dwelling  species,  Monadenia  circum- 
carinata  (Stearns,  1879)  and  Oreohelix  elrodi 
(Pilsbry,  1900)).  A  closer  study  is  needed  to  dis- 
criminate analogous  from  homologous  shell 
character  states  and  extend  the  cladistic  analysis 
to  the  species  level. 

Three  character  complexes,  especially,  illus- 
trate the  problems  left  unsolved  by  this  analysis. 
The  arboreal  habit  in  Monadenia,  s.s.,  is  re- 
garded as  an  apomorphy,  but  many  of  the  as- 


sociated apomorphic  characteristics  (potential 
for  large  size,  high  spire)  are  differentially  ex- 
pressed in  the  various  subspecies  of  the  poly- 
typic Monadenia  fidelis.  Are  these  characters 
secondarily  lost  in  such  subspecies  as  the  low- 
spired  M.  f.  scottiana  Berry  and  the  small- 
shelled,  unnamed  race  near  Klamath  Lake,  or 
do  these  races  preserve  the  plesiomorphous 
condition?  Perhaps  the  arboreal  habit  and  its 
associated  character  states  are  better  viewed  as 
expressions  of  a  physiologically  based  eurytopy 
which  permits  life  in  the  trees  but  also  allows 
M.  fidelis  to  inhabit  low  herbage  near  the  coast 
(M.  f.  pronotis  Berry)  or  rockpiles  (Klamath 
Lake  race).  A  specialized  arboreal  habit,  such 
as  the  dependence  of  M.  setosa  on  standing 
broadleaf  deadwood  for  juvenile  habitat  (Roth 
and  Eng  1980),  is  probably  a  second  level  of 
apomorphy,  derived  from  a  more  generalized 
arboreal  potentiality. 

A  transparent  periostracum,  through  which 
shell  banding  is  visible,  is  probably  plesiomor- 
phous for  the  genus.  Banding,  which  is  wide- 
spread in  Helminthoglyptidae  and  other  helica- 
cean  families,  presumably  evolved  in  the  face  of 
visual  predation  pressure  (Roth,  in  preparation). 
An  opaque  periostracum  that  masks  the  banding 
(still  present  in  the  shell  underneath)  seems  to 
be  associated  with  cryptic  habits  and  indepen- 
dently derived  in  Monadenia,  s.s.,  and  Cory- 
nadenia. 

Carination  of  the  margin  of  the  adult  shell  is 
a  paedomorphy  (compare  Gould  1969,  Poecilo- 
zonites),  independently  derived  in  Monadenia, 
s.s.,  Corynadenia,  and  probably  Shastelix. 
"Hypercarination"  (carina  set  off  by  pinched 
grooves)  is  a  further  derived  state  among  several 
deep  rock-crevice-dwelling  forms  of  the  Sierra 
Nevada. 

While  a  cladogram,  such  as  that  in  Figure  17, 
is  nothing  more  than  a  branching  diagram  de- 
picting the  nested  pattern  of  synapomorphies 
among  the  taxa  under  study,  a  phylogeny  is  an 
explicit  statement  concerning  the  exact  nature 
of  the  evolutionary  relationship  among  the  taxa 
(Eldredge  1979).  A  phylogenetic  tree  is  "a  dia- 
gram (not  necessarily  branching!)  depicting  the 
actual  pattern  of  ancestry  and  descent  among  a 
series  of  taxa"  (Eldredge  1979:168)  and  embod- 
ies assumptions  and  information  not  required  in 
(and  theoretically  excluded  from)  the  construc- 
tion of  a  cladogram.  With  the  further  addition  of 


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401 


explanatory  narrative,  one  can  often  devise  a 
scenario  to  suggest  how  the  phylogenetic  rela- 
tionship, and  the  pattern  of  synapomorphies  in 
the  underlying  cladogram,  came  to  be. 

Strictly  speaking,  the  construction  of  phylo- 
genetic trees  is  limited  to  species;  species,  not 
genera  or  other  higher  taxa,  give  rise  to  other 
species.  Therefore,  when  we  speak  of  a  dichot- 
omy arising  between  subgenera  (as  herein),  we 
are  really  maintaining  that  the  dichotomy  arose 
between  two  species  which  would  be  classed  in 
the  separate  subgenera. 

The  suggested  cladogram  for  the  subgenera  of 
Monadenia  specifies  three  detected  autapomor- 
phies  in  the  subgenus  Monadenia,  s.s.,  two 
synapomorphies  shared  by  Shastelix  and  Cory- 
nadenia,  and  three  autapomorphies  each  in 
Corynadenia  and  Shastelix.  The  effect  of  this 
distribution  of  apomorphies  is  to  falsify  all  pos- 
sible phylogenetic  trees  for  the  group  except  that 
which  is  isomorphous  with  the  cladogram  (see 
Platnick  1977:440-441,  fig.  2).  Consequently,  the 
proposed  phylogenetic  history  of  Monadenia 
consists  of  (1)  a  dichotomy  between  Monaden- 
ia, s.s.,  and  the  common  ancestor  of  Coryna- 
denia and  Shastelix,  followed  in  time  by  (2)  the 
dichotomy  between  Corynadenia  and  Shastelix. 

THE  SCENARIO. — In  the  John  Day  Formation 
(late  Oligocene  to  early  Miocene)  of  central  Or- 
egon, three  fossil  forms  referred  to  Monadenia 
are  present:  M.  antecedens  (Stearns,  1900), 
large-shelled  and  so  similar  in  general  appear- 
ance to  M.  fidelis  that  it  was  formerly  ranked  as 
a  subspecies  of  the  latter;  M.  dubiosa  (Stearns, 
1902),  said  to  be  another  fidelis  -like  form  (Pils- 
bry  1939);  and  M.  marginicola  (Conrad,  1871), 
which  Hanna  (1920)  believed  to  be  related  to 
"the  mormonum  group"  (i.e.,  Corynadenia}. 
No  anatomical  details  are  available,  of  course, 
and  the  shell  microsculpture,  if  preserved,  is  not 
specified  in  the  literature,  but  the  distinction  be- 
tween large  fidelis-lype  shells  and  smaller  shells 
with  the  character  of  present-day  ground-dwell- 
ing forms  is  consistent  with  the  hypothesized 
early  split  between  Monadenia,  s.s.,  and  the 
Shastelix -Corynadenia  stock. 

At  the  time  of  deposition  of  the  John  Day  For- 
mation (36.4-22  million  years  before  present; 
Hammond  1979),  the  Cascade  Arc  had  rotated 
from  its  earlier  northwest-southeast  orientation 
almost  to  its  present  north-south  position,  east 
of  the  presumably  coastal  Klamath  Mountains- 


Coast  Range  block  (Hammond  1979:figs.  8,  9). 
The  John  Day  accumulated  as  clayey  and  tuff- 
aceous  sediments  in  an  inland  basin  contempo- 
raneously with  formation  of  the  volcanic  and 
pyroclastic  rocks  of  the  middle  Western  Cas- 
cades Group.  During  this  episode,  Cascade  vol- 
canism  was  centered  in  randomly  spaced  strato- 
volcanoes  and  calderas;  the  string  of  high  volcanic 
edifices  that  characterizes  the  modern  Cascade 
Range  had  not  yet  developed.  Contemporaneous 
floras  from  the  Coast  Range  block  to  the  west 
are  similar  in  floristic  composition  and  leaf  phys- 
iognomy to  the  Bridge  Creek  Flora  from  the 
lower  member  of  the  John  Day  (Brown  1959; 
Wolfe  and  Hopkins  1967),  indicating  that  the 
Cascade  Range  was  not  a  significant  climatic  or 
vegetational  barrier  at  this  time. 

The  Bridge  Creek  Flora,  associated  with  a  ra- 
diometric  date  of  31.5  million  years  (Evernden 
and  James  1964),  represents  a  mixed  mesophytic 
forest  dominated  by  broad-leaved  deciduous 
trees,  in  a  temperate  climate  with  ample  summer 
rainfall  (Chancy  1948;  Brown  1959;  Wolfe  and 
Hopkins  1967).  Its  greatest  similarities  are  with 
modern  hardwood  forests  of  eastern  North 
America  and  eastern  Asia  (Chaney  1948;  Whit- 
taker  1961).  Temperature  parameters  suitable 
for  forests  of  this  composiion  are  now  lacking 
in  the  Pacific  coast  states  except  in  isolated, 
small,  interior  valleys  (Wolfe  1979). 

The  source  of  the  John  Day  land  mollusks  is 
evidently  the  vertebrate-rich  middle  member,  of 
early  Arikareean  (late  Oligocene)  age  (Wood 
et  al.  1941)  and  about  25  million  years  old 
(Berggren  and  Van  Couvering  1974).  Contem- 
poraneous floras  are  also  mixed  mesophytic, 
possibly  somewhat  warmer  than  the  Bridge 
Creek  Flora.  The  diverse  vertebrate  remains 
likewise  indicate  a  temperate  and  wet  climate, 
quite  unlike  the  semiarid  interior  of  Oregon  of 
the  present.  It  seems  plausible  that  such  an  en- 
vironment could  have  supported  greater  intra- 
generic  snail  diversity  than  now  seen  in  any  for- 
ests of  the  west,  much  as  the  hardwood  forests 
of  the  eastern  United  States  now  support  a  sub- 
stantial diversity  of  snails  in  genera  of  the 
Polygyridae.  Part  of  the  early  diversification  of 
Monadenia  forms  probably  involved  habitat 
partitioning  between  ground-dwelling  and  arbo- 
real species. 

The  subsequent  vegetational  and  climatic  his- 


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tory  of  western  North  America  (greatly  simpli- 
fied) includes  warming  from  late  Oligocene  to 
middle  Miocene,  followed  by  gradual,  fluctuat- 
ing cooling  through  the  Pleistocene,  and  a  shift 
from  summer- wet  to  summer-dry  climate.  By 
the  Pliocene  the  woody  flora  was  depauperate, 
with  extinctions  most  marked  in  broad-leaved 
lineages  with  paratropical  and  subtropical  affin- 
ities (Wolfe  1979).  Latitudinal  and  altitudinal  zo- 
nation  became  pronounced  in  the  later  Tertiary, 
accompanied  by  a  differentiation  of  forest  types 
into  several  adaptive  zones  (Axelrod  1976:fig. 
7).  As  the  rising  Cascade  Range  cast  a  more  pro- 
found rain  shadow  on  the  interior,  humid  forests 
became  confined  to  the  windward,  west  slope  of 
the  Cascades  and  lands  to  the  west — the  modern 
range  of  Monadenia,  s.s. 

At  this  time  (late  Miocene-Pliocene),  a  sub- 
stantial allopatry  between  the  two  existing 
stocks  of  Monadenia  may  have  arisen,  with  the 
Shastelix-Corynadenia  stock  inhabiting  the 
drier,  interior  regions,  and  Monadenia,  s.s., 
perhaps  exploiting  a  tendency  toward  eurytopy 
in  humid  environments.  Equability  of  climate 
may  also  have  been  a  factor.  Parapatry  like  that 
which  now  occurs  in  the  Klamath  Mountains, 
where  M.  (Monadenia)  fidelis  and  M.  (M.)  se- 
tosa  inhabit  riparian  woodland  while  M ' .  (Shas- 
telix) churchi  occupies  drier,  more  exposed 
slopes  and  rockslides,  may  have  been  common 
where  the  environments  interfingered.  (In  this 
connection,  according  to  Whittaker  [1961]  and 
Axelrod  [1976],  the  modern  Klamath  Mountains 
preserve  a  climate  and  vegetation  more  like  that 
of  the  later  Cenozoic  than  any  other  region  in 
the  west.)  Some  time  in  this  interval  Monadenia 
became  extinct  in  central  Oregon. 

On  a  graph  of  mean  annual  temperature  ver- 
sus mean  annual  range  of  temperature  (compare 
Wolfe  1979),  only  the  thermal  range  of  Shastelix 
overlaps  that  of  present-day  mixed  mesophytic 
forest  (Fig.  18).  Because  the  John  Day  fossil  oc- 
currence represents  only  one  datum,  we  do  not 
know  what  the  total  Oligocene  thermal  range  of 
Monadenia  might  have  been;  but  certainly  the 
large,  fidelis-type  snails  of  the  typical  subgenus 
have  shifted  away  (toward  the  right — equable — 
side  of  the  graph)  from  the  thermal  zone  of 
mixed  mesophytic  forest.  Shastelix  has  shifted 
less,  if  at  all.  The  narrow  overlap  of  thermal 
ranges  of  Monadenia,  s.s.,  and  Shastelix  par- 
allels their  narrow  geographic  zone  of  parapatry 


and  suggests  progressive  range/habitat  differ- 
entiation along  a  climatic  gradient.  In  contrast, 
the  thermal  range  of  Corynadenia  overlaps 
those  of  both  other  subgenera  extensively.  It 
occupies  a  considerable  range  of  mean  annual 
temperatures,  but  a  limited  zone  of  mean  annual 
temperature  ranges.  If  Corynadenia  differen- 
tiated from  Shastelix  along  a  climatic  gradient, 
it  was  presumably  not  chiefly  a  gradient  of  mean 
temperature  and  equability. 

The  autapomorphies  of  Corynadenia  can  be 
interpreted  as  adaptations  to  the  rockslide/rock- 
crevice  habitat  and  prolonged  summer  drought. 
The  range  of  Corynadenia  receives  the  least 
precipitation  of  any  zone  inhabited  by  Mona- 
denia (Elford  1970).  Reduction  or  partial  loss  of 
the  genital  apparatus  is  associated  with  adapta- 
tion to  xeric  environments  in  other  groups  of 
Helminthoglyptidae  (Gregg  1960;  Bequaert  and 
Miller  1973).  Retention  of  granulose  microsculp- 
ture  on  the  teleoconch  is  evidently  a  paedomor- 
phy,  and  is  most  strongly  developed  in  species 
(M.  hillebrandi,  M.  circumcarinata)  with  pae- 
domorphic  carination  of  the  adult  shell.  Small 
size  and  low-spired,  lenticular  shape  are  other 
crevice-related  characteristics  that  occur  in 
some  species  and  races  of  Corynadenia. 

Additional  evidence  for  the  relationship  of 
temperature  and  distribution  in  Monadenia  is 
seen  in  the  close  correspondence  between  max- 
imum summer  and  minimum  winter  isotherms 
and  the  range  limits  of  Shastelix  and  Coryna- 
denia (Fig.  1).  The  36  C  mean  maximum  July 
isotherm  practically  coincides  with  the  "down- 
hill" limit  of  the  genus  in  the  Klamath  Moun- 
tains and  Sierra  Nevada,  and  the  -4  C  mean 
minimum  January  isotherm,  more  loosely,  with 
the  "uphill"  limit.  (At  this  scale,  the  isotherms 
are  highly  generalized  and  each  stands  for  a 
range  of  related  microhabitat  temperatures.)  The 
southern  limit  of  Corynadenia  occurs  just  north 
of  where  these  two  isotherms  pinch  together. 
Similarly,  between  the  southern  limit  of  Shas- 
telix and  the  northern  limit  of  Corynadenia,  the 
July  isotherm  bulges  eastward,  almost  but  not 
quite  intersecting  the  January  isotherm.  At  this 
point,  the  33  C  mean  maximum  July  isotherm, 
which  most  closely  coincides  with  the  very 
southern  limit  of  Shastelix,  is  actually  east 
("uphill")  of  the  -4  C  January  isotherm.  If  it 
persisted  long  enough,  a  thermal  configuration 
like  this  could  have  produced  vicariance  within 


ROTH:  DISTRIBUTION,  ANATOMY,  VARIATION  OF  MONADENIA  TROGLODYTES 


403 


20 


15 


10 


Monadenia,  s.  s.x  or 


10  15  20 

Mean  Annual  Range  of  Temp.  (°C) 


25 


30 


FIGURE  18.  Graph  of  thermal  ranges  of  the  three  subgenera  of  Monadenia  in  relation  to  temperature  parameters.  Stippling, 
Monadenia,  sensu  stricto;  diagonal  hachure,  Cory  node  nia;  horizontal  hachure,  Shastelix.  Dot-dashed  line  encloses  thermal 
range  of  present-day  mixed  mesophytic  forest  of  eastern  Asia  (after  Wolfe  1979).  Abbreviations  for  stations  defining  the 
subgeneric  polygons  (in  California  unless  otherwise  indicated):  hh.  Big  Bar;  ca.  Cape  Blanco,  Oreg.;  ch,  Clallam  Bay  1  NNE, 
Wash.;  dn,  Dunsmuir;  du,  Dudley;  er,  Elwha  Ranger  Station,  Wash.;  eu.  Eureka;  gf.  Giant  Forest;  gg,  Grant  Grove;  gr, 
Graton  1  w;  ms.  Mount  Shasta;  or,  Orleans;  pr,  Prince  Rupert,  Brit.  Col.;  .vc,  Scotia;  si,  Sitka,  Alaska;  .vr,  Sonora  Ranger 
Station;  //,  Tatoosh  Island,  Wash.;  vr,  Yreka.  Thermal  data  from  Elford  (1970),  Wolfe  (1979). 


the  Shastelix-Corynadenla  stock.  The  present 
configuration  of  the  isotherms  must  be  short- 
lived in  geologic  terms,  particularly  in  the  face 
of  the  great  climatic  fluctuations  of  the  Pleisto- 
cene; while  it  may  now  enforce  the  geographic 
separation  of  Shastelix  and  Corynadenia,  their 
initial  cleavage  undoubtedly  took  place  earlier. 
Alone,  or  in  combination  with  climatic  factors, 
Miocene  or  later  topographic  developments 
could  have  fostered  the  vicariance.  (Pliocene 
through  Holocene  volcanic  rocks  of  the  High 
Cascades  Group  overlap  the  edges  of  the  north- 
ern Sierra  Nevada  and  eastern  Klamath  Moun- 
tains, separating  the  metamorphic  and  plutonic 
rocks  of  these  two  provinces  for  a  distance  of 
about  80  km  [Hammond  1979].  They  were  ex- 
truded initially  as  basaltic  flows  and  later  as  an- 
desitic,  cone-building  eruptions,  filling  a  struc- 
tural trough  and  lowland  between  the  Klamath 
and  Sierran  blocks.  For  a  calcicolous  ancestral 
Monadenia,  these  volcanic  rocks  could  have 
been  a  barrier  leading  to  allopatric  differentia- 


tion or  interrupting  a  cline  along  which  some 
differentiation  had  already  proceeded.)  Given 
the  complexity  of  relations  between  land  snails 
and  their  environment,  it  is  unlikely  that  a  single 
event  can  be  pinpointed  as  the  cause,  but  no 
details  of  phylogeny,  distribution,  or  thermal  re- 
lations contraindicate  vicariance  in  the  general 
geographic  setting  of  the  present. 

ACKNOWLEDGMENTS 

I  gratefully  acknowledge  the  friendship  and 
guidance  of  Allyn  G.  Smith  (1895-1976),  who 
nourished  my  interest  in  terrestrial  malacology 
and  helped  to  frame  an  early  draft  of  this  study. 
In  addition  I  thank  S.  E.  Hirschfeld,  R.  L.  Seib, 
and  D.  C.  Rudolph,  all  of  whom,  knowing  of  my 
interest  in  Shasta  County  snails,  contributed 
material  for  study.  Special  thanks  are  owed  to 
P.  U.  Rodda,  D.  H.  Kavanaugh,  H.  W.  Schorn, 
P.  H.  Pressley,  and  D.  W.  Taylor  for  helpful 
discussion;  to  A.  J.  Cain,  G.  M.  Davis,  and  W. 
L.  Lee  for  stimulus  in  several  areas;  to  A.  J. 


404 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES.  Vol.  42,  No.  15 


Ferreira  for  radular  preparation;  to  W.  K.  Emer- 
son and  H.  Feinberg  for  lending  historically  sig- 
nificant material;  and  to  L.  L.  Eng,  E.  V.  Coan, 
and  L.  H.  Roth  for  aiding  the  investigation  in 
various  ways.  Susan  Middleton  contributed 
photographic  assistance.  Steven  Sechovec  ren- 
dered most  of  the  anatomical  drawings. 

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northern  California  (abstract).  West.  Soc.  Malacol.  Annu. 
Rep.  11:13. 
,  AND  L.  L.  ENG.    1980.    Distribution,  ecology,  and 


reproductive  anatomy  of  a  rare  land  snail,  Monadenia  se- 

tosa  Talmadge.  Calif.  Fish  Game  66:4-16. 
SMITH,  A.  G.   1957.  Snails  from  California  caves.  Proc.  Calif. 

Acad.  Sci.,  ser.  4,  29:21^6. 
.   1960.  Note  on  Trilobopsis  tehamana  (Pilsbry),  a  rare 

northern  California  land  snail.  Veliger  2:97. 


.    1970.    American  Malacological  Union  Symposium, 

Rare  and  Endangered  Mollusks.  6.   Western  land  snails. 
Malacologia  10:39-46. 

SOLEM,  A.  1975.  Notes  on  Salmon  River  Valley  oreohelicid 
land  snails,  with  description  of  Oreohelix  waltoni.  Veliger 
18:16-30. 

— .  1978.  Classification  of  the  land  Mollusca.  Pp.  49-97 
in  V.  Fretter  and  J.  Peake,  eds.,  Pulmonates,  vol.  2A,  Sys- 
tematics, evolution  and  ecology.  Academic  Press,  London. 

STEARNS,  R.  E.  C.  1900.  Fossil  land  shells  of  the  John  Day 
region  with  notes  on  related  living  species.  Proc.  Wash. 
Acad.  Sci.  2:651-660. 

TALMADGE,  R.  R.  1960.  Color  phases  in  Monadenia  fidelis 
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TWEEDIE,  M.  W.  F.  1961.  On  certain  Mollusca  of  the  Ma- 
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VAN  GOETHEM,  J.  L.  1977.  La  systematique  des  Urocyclinae 
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WAHRHAFTIG,  C.,  AND  J.  H.  BIRMAN.  1965.  The  Quaternary 
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WALTON,  M.  L.  1970.  Longevity  in  Ashmunella,  Monaden- 
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WEBB,  G.  R.  1952.  Pulmonata,  Xanthonycidae:  Comparative 
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WHITTAKER,  R.  H.  1961.  Vegetation  history  of  the  Pacific 
Coast  states  and  the  "central"  significance  of  the  Klamath 
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WOLFE,  J.  A.  1979.  Temperature  parameters  of  humid  to 
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,  AND  D.  M.  HOPKINS.  1967.  Climatic  changes  record- 
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matic changes  in  the  Pacific,  llth  Pac.  Sci.  Congress,  To- 
kyo, 1966,  Sympos.  25. 

WOOD,  H.  E.,  II,  R.  W.  CHANEY,  J.  CLARK,  E.  H.  COLBERT, 
G.  L.  JEPSEN,  J.  B.  REESIDE,  JR.,  AND  C.  STOCK.  1941. 
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406 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  15 


APPENDIX:  LOCALITIES 

Localities  for  material  studied  are  listed  below,  grouped  according  to  geographic  area.  The  num- 
bers assigned  are  for  purposes  of  this  study;  samples  are  on  deposit  in  the  mollusk  collection  of  the 
California  Academy  of  Sciences.  Map  quadrangle  names  refer  to  the  most  recent  edition  of  the 
U.S.  Geological  Survey  7.5-  and  15-minute  Series  (Topographic).  The  number  in  brackets  following 
the  collection  data  is  the  total  number  of  specimens  examined  from  that  locality;  it  does  not  always 
agree  with  sample  size  numbers  cited  in  the  text  because  not  every  character  could  be  measured 
for  every  specimen  and  because  many  of  the  computations  utilized  adult  shells  or  anatomies  only. 
Distances  are  cited  as  received  in  the  collectors'  notes  and  have  not  been  converted  to  the  metric 
system. 


CALIFORNIA:  Sixkiyou  County:  Hornbrook  Quadrangle 

1  Along  creek  1.5  mi  sw  of  Hornbrook.  G  D.  Hanna,  Apr. 
1928  [5]. 

2  SW/4  sec.  18,  T.  46  N,  R.  6  W,  1000  ft  up  Klamath  River 
from  mouth  of  Shasta  River.  D.  E.  Marsh,  1931  [2]. 

3  Among  rocks  about  half-way  up  a  spur  of  Badger  Mountain 
on  west  side  of  Shasta  River  canyon  not  far  above  its 
mouth.  E.  P.  Chace,  29  Sep.  1937  [3].  (Type-locality,  M. 
t.  chuceanu.) 

4  Shasta  River  near  junction  with   Klamath   River.   E.   P. 
Chace  [3]. 

5  Sec.  25(7),  T.  46  N,  R.  7  W,  banks  of  Shasta  River,  2-3 
mi  from  mouth,  in  shaded  rockslides.  A.  G.  Smith,  1 1  Sep. 
1934  [5]. 

CALIFORNIA:  Siskiyou  County:  Yreka  Quadrangle 

6  Chastain's  Quarry,   west  of  Gazelle.    E.   P.   and   E.   M. 
Chace,  28  Sep.  1937  [3]. 

7  [?]  "Near  Yreka."  E.  J.  Elliott,  autumn  1933  [5]. 

CALIFORNIA:     Shasta    County:     Bollibokka     Mountain 
Quadrangle 

8  Near  McCloud  River  bridge,  30  mi  NE  of  Redding.  D.  C. 
Rudolph,  B.  Martin,  S.  Winterath,  9  Apr.  1979  [2]. 

9  Sec.  32,  T.  36  N,  R.  3  W,  west  slope  of  Bollibokka  Moun- 
tain, 0.75  mi  E  of  bridge  across  McCloud  River.  J.  W. 
Durham,  E.  C.  Allison,  18  Apr.  1964  [5]. 

10  Sec.  5,  T.  35  N,  R.  3  W,  slope  below  Samwel  Cave,  above 
McCloud  River  arm  of  Shasta  Lake.  R.  L.  Seib,  1 1  Mar. 
1978  [32]. 

11  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  ch.  1,  sec.  2-5 
(  =  UCMPloc.  1008).  E.  L.  Furlong  [8].  (Type-locality,  M. 
t.  troglodytes.) 

12  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  in  gravel  slope 
filling  grotto  at  south  end,  ch.  2,  sec.  4  (  =  UCMP  loc.  1009). 
E.  L.  Furlong  [2]. 

13  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  with  bone  matter, 
50  ft  in  from  twilight  zone.  R.  E.  Graham,  5-6  Jan.  1957 
[2]. 

14  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  twilight  zone  floor. 
R.  de  Saussure,  10  May  1957  [2]. 

15  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave.  R.  de  Saussure, 
5  June  1957  [2]. 

16  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  entrance  to  main 
cave.  R.  de  Saussure,  5  June  1957  [1]. 

17  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  surface,  second 
exit.  R.  de  Saussure,  A.  Dacey,  5  June  1957  [4]. 

18  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave.  N.  Slusser,  4-7 
June  1957  [2]. 

19  Sec.  5,  T.  35  N,  R.  3  W,  within  25  ft  of  entrance  to  Samwel 
Cave.  R.  E.  Graham,  8  June  1958  [31], 


20  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave,  bone  chamber  pit 
containing  bone  and  rubble  matrix,  7  to  12  inches  depth. 
R.  E.  Graham,  10-14  June  1958  [2]. 

21  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave.  R.  E.  Graham,  27 
Dec.  1958  [6], 

22  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave.  R.  E.  Graham,  18 
June  1959  [1]. 

23  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave.  R.  E.  Graham,  19 
Dec.  1959  [2]. 

24  Sec.  5,  T.  35  N,  R.  3  W,  Samwel  Cave  and  vicinity.  D.  C. 
Rudolph,  B.  Martin,  S.  Winterath,  9  Apr.  1979  [9]. 

25  SE'/a  sec.  7,  T.  35  N,  R.  3  W,  limestone  outcrop,  summit 
of  Hirz  Mountain,  2  mi  sw  of  Samwel  Cave.  R.  E.  Graham, 
June  1960  [2]. 

26  SE'/4  sec.  6,  T.  35  N,  R.  3  W,  Ellery  Creek  on  Oilman 
Road,  15  mi  NE  of  junction  with  Interstate  Hwy.  5,  lime- 
stone rock  slides.  A.  G.  Smith,  17  May  1973:  B.  Roth, 
Mar.  1980  [30]. 

27  SE!4  sec.  6,  T.  35  N,  R.  3  W,  Ellery  Creek,  w  side  of 
McCloud  River  arm  of  Shasta  Lake.  R.  L.  Seib,  24  Mar. 
1978  [5]. 

28  SW/4  sec.  21,  T.  35  N,  R.  3  W,  Dekkas  Rock  at  junction 
of  Dekkas  Creek  with  E  side  of  McCloud  River  arm  of 
Shasta  Lake.  R.  L.  Seib,  24  Mar.  1978  [38]. 

29  NW/4  NE'/4  sec.  32,  T.  35  N,  R.  3  W,  limestone  outcrop 
0.3  mi  ssw  of  junction  of  Campbell  Creek  with  E  side  of 
McCloud  River  arm  of  Shasta  Lake.  R.  L.  Seib,  24  Mar. 
1978  [3], 

30  Sec.  28,  29(7),  T.  25  N,  R.  2  W,  Low  Pass  Creek.  J.  Gor- 
man, 29  Jan.  1953  [2]. 

31  SE'/4  NW/4  sec.  4,  T.  34  N,  R.  2  W,  caves  at  headwaters 
of  Brock  Creek  drainage.  S.  E.  Hirschfeld,  July  1975  [19]. 

32  SE'/4  NW/4  sec.  4,  T.  34  N,  R.  2  W,  shelter  below  and  to 
left  of  Goblin  Shelter,  headwaters  of  Brock  Creek  drain- 
age. S.  E.  Hirschfeld,  July  1975  [8]. 

33  NE'4  SW/4  sec.  3,  T.  34  N,  R.  2  W,  small  cave  in  Hos- 
selkus  limestone  in  gray  rocks  on  E  side  of  eastern  NE- 
sw-trending  ridge.  S.  E.  Hirschfeld,  3  Apr.  1970  [1]. 

34  NW4  NE'4  sec.  8,  T.  34  N,  R.  2  W,  cave  between  two 
limestone  buttes  at  south  end  of  Gray  Rocks,  above  Pit 
River  arm  of  Shasta  Lake.  S.  E.  Hirschfeld,  Oct.  1975  [49]. 
(Type-locality,  M.  t.  wintu.) 

35  SW/4  NW/4  sec.  7,  T.  34  N,  R.  2  W,  limestone  quarry  1 
mi  N,  s/s  mi  E  of  Brock  Mountain  Lookout,  above  Squaw 
Creek  arm  of  Shasta  Lake.  R.  L.  Seib,  25  Mar.  1978  [39]. 

36  Squaw  Creek.  A.  M.  Strong,  1898  [3]. 

37  Squaw  Creek.  J.  Gorman,  14-15  Mar.  1953  [5]. 

38  N  ctr.,  SE'/4  sec.  9,  T.  34  N,  R.  2  W,  caves  in  Brock  Creek 
limestone,  above  Pit  River  arm  of  Shasta  Lake.   S.   E. 
Hirschfeld,  July  1975  [7]. 


ROTH:   DISTRIBUTION,  ANATOMY,  VARIATION  OF  MONADENIA  TROGLODYTES 


407 


39  NW/4  SE!4  sec.  9,  T.  34  N,  R.  2  W,  lower  limestone  across 
from  Brock  Spring,  above  Brock  Creek,  above  Pit  River 
arm  of  Shasta  Lake.  S.  E.  Hirschfeld,  July  1975  [13]. 

40  NE!4  SW'4  sec.  9,  T.  34  N,  R.  2  W,  "Monadenia  Cave" 
and  vicinity,  above  Pit  River  arm  of  Shasta  Lake.  S.  E. 
Hirschfeld,  July  1975  [14]. 

41  NE'4  SW/4  sec.  9,  T.  34  N,  R.  2  W,  "Elk  Antler  Cave," 
above  Pit  River  arm  of  Shasta  Lake.  S.  E.  Hirschfeld,  July 
1975  [20]. 

CALIFORNIA:  Shasta  County:  Lamoine  Quadrangle 

42  SW/i  sec.  13,  T.  34  N,  R.  4  W,  Shasta  Lake  Caverns.  D. 
C.  Rudolph,  B.  Martin,  S.  Winterath,  10  Apr.  1979  [1]. 

43  SE'/i  SE!4  sec.  23,  T.  34  N,  R.  4  W,  Potter  Creek,  along 
McCloud  River  arm  of  Shasta  Lake.  R.  L.  Seib,  11  Mar. 
1978  [22]. 


44  Sec.  23,  T.  34  N,  R.  4  W,  Potter  Creek  Cave,  "past  kitch- 
en" (  =  UCMPIoc.  1055)  [12]. 

CALIFORNIA:  Shasta  County:  Project  City  Quadrangle 

45  SE'4  sec.  4,  T.  33  N,  R.  4  W,  Calveris  Cement  Co.  quarry 
above  Interstate  Hwy.  5  just  south  of  Shasta  Lake  bridge, 
2.5  mi  NNE  of  Mountain  Gate.  R.  L.  Seib,  11  Mar.  1978 
[40]. 

CALIFORNIA:  Shasta  County:  Anderson  Quadrangle 

46  Clear  Creek,  on  road  between  Redding  and  Beegum.  G  D. 
Hanna,  Sep.  1952  [1]. 

CALIFORNIA:  Shasta  County:  quadrangle  uncertain 

47  Crystal  Shasta  Cave.  K.  Howard,  1958(?)  [1]. 

48  Chute  Cave.  R.  de  Saussure,  Nov.  1957  [1]. 

49  "Shasta  County,  Calif.,"  ex  D.  O.  Mills  collection,  UCMP 

[3]. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  16,  pp.  409^433,  2  figs.,  9  tables  October  26,  1981 


FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 
FROM  NORTHERN  CALIFORNIA 

By 
Robert  E.  Jones 

Museum  of  Vertebrate  Zoology,  University  of  California, 
Berkeley,  California  94720 


ABSTRACT:  General  food  habits  are  discussed  for  11  species  of  small  marine  mammals  beach-cast  in  northern 
California.  The  collection  method  allowed  tabulation  only  of  the  kinds,  numbers,  and  percentages  of  fish, 
molluscan,  and  crustacean  prey. 

Dominant  food  species  for  all  male  Zalophus  californianus  were  the  following:  Merluccius  productus  (62.8 
percent  of  occurrence),  Engraulis  mordax  (23.8  percent),  Scorpaenidae  (6.3  percent),  Porichthys  notatus  (3.2 
percent).  Octopus  and  Loligo  also  occurred  in  7  (23.0  percent)  of  30  California  sea  lions  examined. 

Nine  of  19  individuals  of  Eumetopias  jubatus  contained  food  remnants  of  fish;  10  new  dietary  items  are 
reported.  Seven  of  the  nine  also  fed  on  invertebrates,  including  four  kinds  of  cephalopods.  Scorpaenidae  31.2 
percent  of  occurrence),  Merluccius  productus  (21.7  percent),  Pleuronectidae  (17.3  percent),  and  Chilara  taylori 
(11.8  percent)  were  the  principal  fishes  identified. 

Eight  of  12  Phoca  vitulina  stomachs  contained  food  (6  with  fish,  2  with  invertebrates).  Embiotocids  (41.9 
percent  of  occurrence),  Lycodopsis  padfica  (27.9  percent),  Pleuronectidae  (9.3  percent),  and  Hexagrammos 
decagrammus  (9.3  percent)  were  the  dominant  fishes.  The  only  cephalopod  identified  from  Phoca  was  Octopus. 

The  stomachs  of  all  seven  individuals  of  Lagenorhynchus  obliquidens  examined  contained  food.  Osmerid 
fishes,  Porichthys,  and  juvenile  rockfishes  were  the  most  frequent  items  identified.  The  presence  of  five  genera 
of  cephalopods  suggests  that  Lagenorhynchus  can  take  a  wide  variety  of  prey  from  several  habitats. 

Juvenile  Scorpaenidae,  Engraulis,  Merluccius,  and  Microgadus  made  up  97  percent  of  the  diet  of  20  individuals 
ofPhocoena  phocoena.  The  abundant  cephalopod  Loligo  was  a  primary  invertebrate  food  item  and  was  ingested 
whole. 

INTRODUCTION 

The  objective  of  this  study  is  to  summarize  mercial  fishing  industries  (Anonymous    1901; 

the  literature  on  the  diets  of  smaller  marine  Townsend  1918). 

mammals  in  the  eastern  Pacific  and  to  report  on          Marine  biologists  in  California  have  ques- 

the  contents  of  102  stomachs  from  five  pinniped  tioned  the  impact  of  marine  mammals  on  the 

and  six  small  cetacean  species.  The  purpose  has  state's  fisheries  for  at  least  100  years  (Redding 

been  to  make  a  thorough  scrutiny  of  published  etal.  1875;  Rutteretal.  1904;  California  Division 

dietary  knowledge  and  analyze  stomachs  of  of  Fish  and  Game  1927;  Hedgpeth  1944;  Ander- 

beach-cast  specimens.  Particular  attention  has  son  1960;  Frey  1971;  National  Oceanic  and  At- 

been  directed  to  reviewing  all  records  on  feeding  mospheric  Administration  1974).  The  report  of 

controversy.  It  has  been  argued  for  many  de-  the  California  Commissioners  of  Fisheries  for 

cades  that  marine  mammals  compete  with  com-  the  years  1874-1875  (Redding  et  al.  1875)  stated 

[409] 


410 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


clearly  that  sea  lions  and  seals  were  protected 
by  special  enactment,  with  penalties  imposed  on 
any  person  who  should  kill  or  disturb  the  beasts. 
On  the  basis  of  the  observed  "hundred-fold" 
increase  in  the  populations  of  seals  and  sea  lions 
at  the  entrance  to  San  Francisco  Bay  over  the 
previous  ten-year  period,  the  Commissioners 
recommended  the  killing  of  nine-tenths  of  the 
existing  population.  It  is  unclear  as  to  what  ac- 
tion, if  any,  resulted  from  this  recommendation. 
Scammon  (1874)  stated,  "A  few  years  ago  great 
numbers  of  sea  lions  were  taken  along  the  coast 
of  Upper  and  Lower  California,  and  thousands 
of  barrels  of  their  oil  were  obtained."  The  state- 
ment describes  the  period  before  the  enactment 
of  regulations  protecting  California  pinnipeds. 
On  the  west  coast  of  North  America,  most  of 
the  commercial  seal  and  sea-lion  killing  stopped 
about  1860  (Bonnot  1928). 

The  problem  of  a  declining  fishery  was  aired 
before  another  commission  at  San  Francisco  in 
1899,  and  on  27  April  1899,  permission  was 
granted  to  kill  sea  lions  along  the  coast,  includ- 
ing lighthouse  reservations  (Bonnot  1928).  The 
data  from  stomach  inspections,  gathered  by  Pro- 
fessor L.  L.  Dyche  in  1899,  were  published  by 
C.  H.  Merriam  (1901a,  1901b).  These  publica- 
tions were  an  attempt  toward  a  critical  exami- 
nation of  stomachs  from  slaughtered  sea  lions. 
A  view  held  by  many  was  that  stomach  analysis 
was  utterly  useless,  for  the  observed  fact  was 
that  sea  lions  pursued  salmon  through  the  Gold- 
en Gate  as  far  as  Sacramento  (Anonymous 
1901).  Thousands  were  killed  before  the  Treas- 
ury Department  prohibited  wholesale  slaughter 
on  government  lands  on  31  May  1899  (Merriam 
1901a).  The  destruction  of  sea  lions  was  justified 
by  the  belief  that  the  declining  shad  (Alosa  sap- 
idissima),  salmon  (Oncorhynchus  spp.),  and 
striped  bass  (Morone  scixatilis)  fisheries  would 
thereby  increase,  but  little  evidence  was  gath- 
ered to  verify  or  deny  such  claims.  The  U.S. 
Fish  Commission,  the  Secretary  of  the  U.S.  De- 
partment of  Agriculture,  and  the  New  York 
Zoological  Society  opposed  the  decisions  of  the 
California  Fish  Commission  (Anonymous 
["Steelhead"  pseudonym]  1901). 

The  controversy  over  control  of  sea  lion  pop- 
ulations pitted  biologists  C.  Hart  Merriam,  Bar- 
ton W.  Evermann,  and  William  T.  Hornaday 
against  David  Starr  Jordan,  Charles  H.  Gilbert, 
Harvey  W.  Harkness,  and  N.  Baird  Scofield 


(Starks  1918;  Townsend  1918,  1919).  The  1904 
report  of  a  Federal  commission  appointed  to  re- 
view the  situation  contains  the  first  published 
list  of  stomach  contents  of  a  series  of  California 
sea  lions  (Zalophus)  and  northern  sea  lions  (Eu- 
metopias)  (Rutter  et  al.  1904). 

In  1914-1916,  the  Newcombe  Commission 
gathered  basic  biological  field  data  in  British 
Columbia  (Newcombe  and  Newcombe  1914; 
Newcombe  et  al.  1918).  Twelve  of  the  14  north- 
ern sea  lions  examined  contained  intact  herring; 
however,  food  items  consumed  by  these  sea 
lions  throughout  the  year  were  not  documented. 

Early  in  the  20th  century,  suggestions  were 
made  to  harvest  young  sea  lions,  following  the 
example  of  the  Newfoundland  harp-seal  indus- 
try. Harp  seals  (Phoca}  had  been  cropped  an- 
nually for  100  years  with  an  annual  take  of  pups 
exceeding  200,000.  Recommendations  for  con- 
trol of  pinnipeds  in  California  gained  the  support 
of  many  prominent  biologists  (Townsend  1919). 

Following  an  investigation  at  the  lighthouse 
reservation  on  Ano  Nuevo  Island  (Evermann 
1921),  it  was  stated  that  surplus  bulls  on  this 
rookery  could  be  killed  (Evermann  and  Hanna 
1925).  This  is  the  earliest  example  of  a  recom- 
mendation to  harvest  marine  mammals  along  the 
California  coast  based  on  biological  information. 
Because  sea  lions  and  seals  are  polygamous,  it 
was  felt  that  10  percent  of  the  excess  males 
could  be  killed,  but  for  economic  reasons  the 
cropping  was  not  carried  out  (Rowley  1929). 
Target  clubs  obtained  military  surplus  guns  and 
ammunition  and  "practiced"  on  the  Santa  Cruz 
and  San  Mateo  coast  pinniped  populations.  Ap- 
parently this  activity  frightened  the  pinnipeds 
enough  to  cause  their  departure  (Herb  Steindorf, 
local  rancher,  pers.  comm.,  1969).  Fishermen 
still  ask  permission  to  kill  sea  lions,  and  numer- 
ous reports  of  damage  to  the  fishery  continue  to 
this  day. 

Between  the  late  1920's  and  1972,  when  the 
Marine  Mammal  Protection  Act  was  passed, 
certain  marine  mammals  were  fully  protected, 
but  commercial  and  sport  fishermen  (actively 
fishing  from  boats)  could  kill  sea  lions  and  har- 
bor seals  interfering  with  their  operations  (Cal- 
ifornia Fish  and  Game  Code,  Sections  3002, 
4500).  Thus  fishermen  could  protect  their  nets, 
tackle,  and  fish  from  damage  by  marine  mam- 
mals. The  California  Fish  and  Game  Commis- 
sion reserved  the  right  to  reduce  the  population 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


411 


and  to  require  permits  for  educational  display  or 
scientific  taking  of  sea  lions  or  seals.  The  sea 
otter  (Enhydra  lutris),  Guadalupe  fur  seal  (Arc- 
tocephalus  townsendi),  and  elephant  seal  (Mi- 
rounga  angustirostris)  were  fully  protected. 
State  laws  applied  only  to  the  seals,  sea  lions, 
and  sea  otter;  no  regulations  concerning  ceta- 
ceans were  in  effect  until  the  enactment  of  the 
Marine  Mammal  Protection  Act  of  1972.  This 
protection  placed  an  immediate  moratorium 
upon  the  taking  and  importation  of  all  marine- 
mammal  products  into  the  United  States  (Na- 
tional Oceanic  and  Atmospheric  Administration 
1974).  The  oft-repeated  complaint  about  the  lack 
of  life  history  data  on  most  of  our  common 
species  of  marine  mammals  is  still  valid  (Starks 
1918;  Anderson  1960;  Peterson  and  Bartholo- 
mew 1967;  Briggs  and  Davis  1972).  Even  the  age 
at  puberty  and  physical  maturity  of  the  Califor- 
nia sea  lion  (Zalophus  californianus)  is  un- 
known (Harrison  1972).  There  are  few  published 
accounts  of  marine  mammal  life  histories,  and 
these  studies  seldom  contain  precise  information 
on  food  habits  (Bartholomew  1967;  Orr  and 
Poulter  1965,  1967;  Peterson  and  Bartholomew 
1967;  Peterson  and  LeBoeuf  1969;  Odell  1971; 
Seed  1972).  General  publications  on  marine 
mammals  seldom  contain  significant  or  specific 
information  on  food  habits  (Sergeant  and  Fisher 
1957;  Slijper  1962;  King  1964;  Evans  and  Bastian 
1969;  Daugherty  1972;  Orr  1972;  Ridgway  1972). 

Notes  on  food  habits  of  pinnipeds  have  been 
reported  in  the  literature  since  the  early  work  of 
L.  L.  Dyche  in  1903  (Bonnot  1928,  1932a,  1932b, 
1951;  Scheffer  and  Neff  1948;  Scheffer  1950a; 
Mathisen  1959;  Mathisen  et  al.  1962;  Thorstein- 
son  and  Lensink  1962;  Fiscus  and  Baines  1966; 
Morejohn  and  Baltz  1970;  Briggs  and,  Davis 
1972). 

The  literature  on  distribution,  food  habits,  and 
life  histories  of  small  cetaceans  is  less  extensive, 
and  wholly  inadequate.  While  the  federal  status 
report  to  the  Secretary  of  Commerce  (National 
Oceanic  and  Atmospheric  Administration  1974) 
is  the  most  complete  summary  of  studies  to  date, 
natural  history  data  are  lacking  even  for  the 
most  common  species.  Cetaceans  have  not  had 
a  reputation  for  eating  commercially  important 
fishes  nor  for  harassing  fishermen.  Until  1972 
there  were  no  federal  laws  protecting  or  regu- 
lating the  capture  of  small  whales  or  porpoises. 
Recently  large  numbers  of  porpoises  have  been 


killed  during  tuna-fishing  operations  (Perrin 
1970).  Papers  dealing  with  food  habits  of  ceta- 
ceans are  widely  scattered  (Scheffer  1950b, 
1953;  Brown  and  Norris  1956;  Tomilin  1957; 
Wilke  and  Kenyon  1952,  1957;  Wilke  and  Ni- 
cholson 1958;  Norris  and  Prescott  1961;  Fitch 
and  Brownell  1968;  Loeb  1972;  Perrin  et  al. 
1973). 

More  study  is  needed  on  feeding  rates  and 
feeding  phenomena  (Sergeant  1968,  1969).  Fast- 
ing periods  and  basic  metabolic  rates  obviously 
affect  calculations  of  the  impact  of  marine  mam- 
mals on  the  food  resources  of  the  sea.  The  role 
of  marine  mammals  in  overall  ocean  ecology 
needs  further  study.  Current  knowledge  of  food 
chains  and  trophic  relationships  of  marine  mam- 
mals has  only  recently  been  given  attention  by 
marine  ecologists  (Steele  1970). 

MATERIALS  AND  METHODS 

During  this  study,  I  examined  102  specimens 
found  dead  on  California  beaches  (Table  1).  The 
11  species  of  marine  mammals  studied  (Fig- 
ures 1  and  2)  include  two  phocids,  Mirounga 
angustirostris  and  Phoca  vitulina,  and  three 
otariids,  Eumetopias  jubatus,  Zalophus  califor- 
nianus, and  Callorhinus  ursinus.  The  six  ceta- 
cean species  represent  three  families:  Phocoe- 
nidae,  Phocoena  phocoena  and  Phocoenoides 
dalli;  Delphinidae,  Delphinus  delphis,  Grampus 
griseus,  and  Lagenorhynchus  obliquidens',  and 
Physeteridae,  Kogia  simus. 

Where  feasible,  specimens  up  to  about  100 
pounds  (45  kg)  were  removed  to  the  laboratory 
and  data  were  recorded  on  standardized  sheets 
(Norris  1961;  Scheffer  1967).  Where  terrain  or 
tidal  conditions  did  not  allow  removal,  carcasses 
were  measured  in  place.  All  possible  standard 
measurements  were  taken. 

The  skull,  baculum  (when  present),  and  stom- 
ach were  collected  from  specimens  too  large  to 
move.  All  material  was  labeled  and  taken  to  the 
laboratory  for  careful  examination.  Osteological 
material  from  each  specimen  was  tagged  and 
cleaned  by  standard  museum  techniques.  Thir- 
teen stomachs  were  donated  by  personnel  of  the 
California  Academy  of  Sciences. 

After  initial  external  cleaning,  25  stomachs 
were  filled  with  cool  tap  water  and  measured  to 
obtain  an  average  stomach  volume  (Table  2). 
Each  water-filled  stomach  was  drained  of  its 


412 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


TABLE  1.     NUMBER  OF  MARINE  MAMMAL  STOMACHS  EXAMINED  IN  NORTH-CENTRAL  CALIFORNIA,  1968-1973;  (n  =  102). 
A  total  of  68  stomachs  contained  food  items;  of  these,  61  had  fish  remains,  33  invertebrates. 


CETACEANS 


Phocoe-      Lagenorhyn- 
Phocoenu          noitles  chus  Delphinus        Grampus  Kogia 


Total 


Examined 

20 

5 

7 

1 

1                        1 

35 

With  contents 

16 

2 

7 

1 

1                        1 

28 

Empty 

4 

3 

0 

0 

0                     0 

7 

With  fish 

14 

2 

5 

1 

0                      1 

23 

With  invertebrates 

8 

0 

3 

1 

1                1 

14 

PINNIPEDS 

Ztilophus 

Euinetopias 

Phoca 

Callorhinus 

Mirounga 

Total 

Examined 

30 

19 

12 

4 

-> 

67 

With  contents 

20 

9 

8 

1 

-> 

40 

Empty 

10 

10 

4 

3 

0 

27 

With  fish 

20 

9 

6 

1 

2 

38 

With  invertebrates 

7 

7 

•> 

-> 

0 

19 

With  rock  or  plant  material 

1 

3 

0 

0 

0 

4 

contents  into  a  graduated  cylinder  for  accurate 
volumetric  reading. 

Stomach  contents  were  segregated,  using  a 
parasite-recovery  technique  of  flotation  and  de- 
canting, and  then  washed  repeatedly  with  clean 
water.  This  method  of  sedimentation-decanta- 
tion  allows  speedy  recovery  of  all  items.  Small 
otoliths  (lapilli)  were  occasionally  detected,  but 
all  otoliths  reported  here  were  sagittae.  Other 
workers  (Fitch  and  Brownell  1968;  Morejohn 
and  Baltz  1970;  Smith  and  Gaskin  1974)  have 
screened  gastro-intestinal  materials  through 
cheesecloth  gauze  or  graded  wire  screens. 

Disarticulated  bones  were  separated  from  in- 
ternal parasites  with  forceps.  Fish  bones  and 
otoliths  were  dried  overnight.  Soft  tissues  were 
preserved  in  alcohol. 

The  heavy  fish  otoliths  (sagittae)  were  easily 
seen  by  moving  the  clear  glass  container  over 
backgrounds  of  several  different  colors.  Even 
the  tiny  otoliths  (asterisci  or  lapilli)  could  be  de- 
tected when  they  were  gently  moved  by  the  cov- 
ering water.  I  removed  the  otoliths  with  a  camel 
hair  brush  or  fine  forceps.  They  were  carefully 
cleaned  of  mucus  (film),  dried,  and  stored  in  gel- 
atin pill  containers.  John  E.  Fitch,  California 
Department  of  Fish  and  Game,  provided  iden- 
tifications. 

Most  cephalopod  beaks,  which  were  stored 
in  alcohol,  were  identified  using  the  pictorial 
guide  developed  by  the  California  Department 
of  Fish  and  Game  (Iverson  and  Pinkas  1971). 
Several  types  were  identified  by  Clifford  Fiscus, 
National  Marine  Mammal  Laboratories,  U.S. 


Department  of  Commerce.  I  later  checked  these 
samples  using  the  keys  developed  by  Clarke 
(1962)  and  by  comparing  them  with  squid  beaks 
furnished  by  Jerome  L.  Spratt  of  the  California 
Department  of  Fish  and  Game.  Four  hundred 
thirty-five  cephalopod  beaks  and  2828  fish 
otoliths  were  identified.  Thirty-one  otoliths  and 
217  cephalopod  beaks  were  not  identifiable.  I 
was  able  to  identify  most  cephalopod  beaks  only 
to  genus. 

Few  stomachs  contained  intact  fish.  Fish  skel- 
etal material  was  air-dried,  and  its  volume  was 
measured  in  a  graduated  cylinder  (Table  9).  Fish 
remains  were  identified  with  the  aid  of  Clothier 
1950;  Roedel  1953;  Clemens  and  Wilby  1961; 
Fitch  and  Lavenberg  1971;  and  Miller  and  Lea 
1972.  Common  names  used  follow  those  of  the 
American  Fisheries  Society  (Robins  et  al.  1980) 
except  for  Atka-mackerel  and  jackmackerel, 
which  follow  Hubbs,  Follett,  and  Dempster 
(1979).  All  recovered  materials  (specimens  and 
contents)  are  deposited  in  the  Museum  of  Ver- 
tebrate Zoology,  Berkeley,  or  the  California 
Academy  of  Sciences,  San  Francisco. 

A  total  of  35  cetacean  and  67  pinniped  car- 
casses was  beach-cast  along  north-central  Cal- 
ifornia. Voucher  material  has  been  collected  and 
deposited  at  these  institutions:  CAS,  California 
Academy  of  Sciences,  San  Francisco;  MVZ, 
Museum  of  Vertebrate  Zoology,  University  of 
California,  Berkeley;  HSC,  Humboldt  State 
University,  Vertebrate  Zoology  Museum,  Ar- 
cata;  PORE,  Point  Reyes  National  Seashore, 
Point  Reyes. 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


413 


FIGURE  1.     Localities  of  cetacean  beach-cast  specimens  in  north-coastal  California,  1968-1973. 


CETACEAN 

Humboldt  Co.—Phocoemi:  HSC  68-7,  F,  2  Sep.  1968,  122 
cm.  12  otoliths;  HSC  73-4,  M,  21  Aug.  1973,  150  cm  est.,  14 
otoliths.— Lagenorhynchus:  HSC  68-9,  M,  26  Dec.  1968,  179 
cm,  36  otoliths,  1  cephalopod  beak. 

Sonoma  Co.—Phocoenoules:  MVZ  153258  (REJ  670),  F,  28 
June  1973,  102  cm,  empty. 

Marin  Co.—Phocoena:  CAS  16602  (2385),  M,  1  June  1973, 
138  cm,  1083  otoliths;  CAS  16603  (2385),  I  June  1973,  146  cm, 
empty;  CAS  21380  (REJ  448),  F,  3  July  1970,  158  cm,  15 
otoliths.— Phocoenoiiles:  MVZ  153259  (REJ  678),  F,  7  Aug. 
1973,  186  cm,  empty;  CAS  16604  (2385),  M,  1  June  1973,  198 
cm,  empty;  CAS  16297  (2335),  M,  21  Sep.  1972,  188  cm,  6 
oloXMis.—Lagenorhynchus:  MVZ  140845  (REJ  218),  F,  9  Feb. 

1970,  181  cm,  no  ID.—Delphinur.  CAS   16242  (2340),  F,  6 
Nov.  1972,  168  cm,  II  otoliths,  4  cephalopod  beaks. 

San  Francisco  Co.—Phocoena:  CAS  16629  (2384),  M,  26 
May  1973,  140  cm,  2  otoliths;  CAS  16572  (2398),  F,  20  Aug. 
1973,  137cm,  17  otoliths.— Grampus:  MVZ  153257  (REJ  659), 
F,  20  May  1973,  275  cm,  barnacle  and  hydroid. 

San  Mateo  Co.—Phocoena:  CAS  15992  (2237),  F,  19  July 

1971,  143  cm,  33  otoliths;  CAS  16609(2390),  F,  13  July  1973, 
159  cm,  26  otoliths  and  1  hake;  CAS  16633  (2392),  F,  13  July 
1973,   126  cm,  4  otoliths,  35  pair  cephalopod  beaks  and   13 
whole  Loligo.—Phocoenoides:  CAS  (REJ  674),  F,   1   Aug. 


1973,  (79  in.),  223  cm,*  61  otoliths.— Lagenorhynchus:  CAS 
16593  (2336),  M,  28  Sep.  1972,  190  cm,  47  otoliths;  CAS  16342 
(2380),  F,  20  Apr.  1973,  193  cm,*  4  otoliths.—  KoKia:  CAS 
16635  (2382),  M,  25  May  1973,  204  cm,  2  otoliths  and  231 
cephalopod  beaks. 

Santa  Cruz  Co.—Phocoena:  CAS  21381  (KB  19-73),  M,  13 
Apr.  1973,  134  cm,  17  otoliths;  (KB  17-73),  F,  1973,  104  cm, 
empty. 

Monterey  Co.—Phocoena:  CAS  21387  (REJ  687),  F,  22 
Sep.  1973,  107  cm,  empty;  CAS  21383  (REJ  673),  M,  24  July 
1973,  126  cm,  48  otoliths,  1  pair  cephalopod  beaks;  CAS  21385 
(REJ  661),  F,  17  June  1973,  104  cm,  16  otoliths.  3  cephalopod 
beaks;  CAS  21386  (REJ  654),  F,  24  Apr.  1973,  137  cm,  15 
otoliths;  CAS  21389  (REJ  653),  M,  25  Apr.  1973,  145  cm,  13 
cephalopod  beaks;  CAS  21388  (REJ  450),  ?  sex,  24  July  1971, 
no  tl,  127  otoliths,  52  cephalopod  beaks;  CAS  21384  (REJ 
449),  ?  sex,  24  July  1971,  108  cm,  empty;  CAS  21382  (REJ 
241),  F,  6  June  1970,  172  cm,  16  cephalopod  beaks.— Lage- 
norhynchus:  CAS  21370  (REJ  652),  F,  21  Apr.  1973,  187  cm, 
4  otoliths;  CAS  21378  (REJ  625),  M,  14  May  1973,  177  cm, 
147  cephalopod  beaks;  MVZ  (REJ  237),  F,  29  May  1970,  180 
cm,  7  cephalopod  beaks. 


Indicates  original  data  taken  in  inches. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


O   Zalophus 

Eumetopias 
•   Phoca 
A   Callorhinus 


FIGURE  2.     Localities  of  pinniped  beach-cast  specimens  in  north-coastal  California,  1968-1973. 


PINNIPEDS 

Humboldt  Co.— Zalophus:  CAS  21401  (REJ  248),  M,  16 
June  1970,  256  cm,  17  otoliths;  HSC  3206  (69-34),  M,  1  June 
1969,  233  cm,*  52  otoliths;  HSC  3205  (REJ  69-33)  M,  27  May 

1969,  228  cm,*   122  otoliths;  HSC  3204  (REJ  69-32),  M,  23 
May  1969,  249  cm,*  93  otoliths;  HSC  3125  (REJ  69-31),  M,  14 
May  1969,  239  cm,*  152  otoliths.— Eumetopias:  CAS  21391 
(REJ  683),  F,  16  Sep.  1973,  1 1 1  cm,  empty;  CAS  21390  (REJ 
672),  M,  20  July  1973,  309  cm,  empty;  HSC,  (REJ  655),  F,  1 
May  1973,  200  cm,  stones  only;  HSC,  (REJ  247),  M,  14  June 

1970,  257  cm,  76  otoliths,  2  cephalopod  beaks;  MVZ  140847 
(REJ  249),  F,  16  June  1970,  259  cm,  66  otoliths,  13  cephalopod 
beaks;  HSC,  (REJ  69-8),  F,  16  Feb.  1969,  251  cm,*  2  otoliths, 
1  cephalopod  beak.— Phoca:  CAS  21421  (REJ  684),  F,  17  Sep. 
1973,  147  cm,  empty;  HSC  1 188  (REJ  69-24),  F,  25  Apr.  1969, 
149  cm,  13  otoliths;  HSC,  (REJ  68-38),  M,  28  July  1968,  162 
cm,  3  otoliths;  HSC,  (REJ  68-14),  F,  6  Apr.  1968,  102  cm,  17 
otoliths.— Mirounga:  HSC  2165  (REJ  69-6),  F,  19  Feb.  1969, 
152  cm,  1  Apristurus  brunneus  egg;  HSC  1356  (?),  F,  5  May 
1970,  no  tl,  26  otoliths. 

Sonoma  Co.— Zalophus:  CAS  21402  (REJ  680),  M,  17  Aug. 
1973,  238  cm,  empty;  CAS  21403  (REJ  660),  M,  5  June  1973, 
215  cm,  98  otoliths,  1  cephalopod  beak. — Eumetopias:  CAS 

21392  (REJ  688),  M,  23  Sep.   1973,  152  cm,  4  otoliths;  CAS 

21393  (REJ  679),  F,  17  Aug.  1973,  229  cm,  empty. 


San  Francisco  Co..— Callorhinus:  MVZ  140846  (REJ  212), 
F,  5  Feb.  1970,  131  cm,  7  isopods. 

Marin  Co.— Zalophus:  CAS  21404  (REJ  700),  M,  29  Sep. 
1973,  210  cm,  empty;  PORE  138  (REJ  657),  M,  9  May  1973, 
230  cm,  I  otolith;  CAS  21405  (REJ  641),  M,  1  Mar.  1973,  220 
cm,  empty;  CAS  16184  (2316),  M,  19  July  1973,  158  cm,  I 
otolith,  1  cephalopod.— Eumetopias:  CAS  21399  (REJ  682), 
M,  6  Sep.  1973,  325  cm,  empty;  CAS  21395  (REJ  677),  F,  7 
Aug.  1973,  235  cm,  broken  otoliths  and  rocks;  PORE  136  (REJ 
676),  M,  6  Aug.  1973,  280  cm,  empty;  CAS  21395  (REJ  668), 
F,  22  June  1973,  234  cm,  empty;  ?(REJ  637),  21  Oct.  1972,  no 
tl,  22  otoliths,  4  cephalopod  beaks;  CAS  21397  (REJ  635),  F, 
16  Sep.  1972,  232  cm,  empty;  CAS  213%  (REJ  629),  M,  22 
July  1972,  228  cm,  59  otoliths,  5  cephalopod  beaks;  PORE  137 
(REJ  453),  F,  14  Aug.  1971,  226  cm,  empty.— Phoca:  PORE 
214  (REJ  681),  F,  29  Aug.  1973,  140  cm,  empty;  CAS  21423 
(REJ  669),  M,  22  June  1973,  97  cm,  3  Crago  sp.;  CAS  21422 
(REJ  642),  M,  1  Mar.  1973,  158  cm,  empty. 

San  Mateo  Co.— Zalophus:  CAS  21412  (REJ  698),  M,  28 
Sep.  1973,  160  cm,  empty;  CAS  2141 1  (REJ  697),  M,  28  Sep. 
1973,  no  tl,  empty;  CAS  21410  (REJ  695),  M,  28  Sep.  1973, 
218  cm,  empty;  CAS  21409  (REJ  693),  M,  28  Sep.  1973,  213 
cm,*  empty;  ?(REJ  692),  M,  28  Sep.  1973,  152  cm  (est.),  2 
otoliths,  2  cephalopod  beaks;  CAS  21408  (REJ  691),  F,  28 
Sep.  1973,  no  tl,  empty;  CAS  21407  (REJ  690),  M,  28  Sep. 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS  415 

TABLE  2.     AVERAGE  STOMACH  VOLUME  OF  THREE  SPECIES  OF  MARINE  MAMMALS  AS  MEASURED  BY  WATER  DISPLACEMENT. 


Total  length  of  animal  (cm) 

Volume  of  stomachs  (  1) 

Species 

n 

Sex 

Range 

Average 

Range 

Average 

Zalophus 

Adults 

6 

M 

198-256 

223 

9.42-19.84 

12.63 

Subadults 

4 

M 

126-160 

145 

1.72-9.12 

5.75 

Eumetopias 

3 

M 

280-325 

305 

22.20-46.72 

33.67 

4 

F 

200-235 

230 

9.80-23.74 

17.78 

Phocoena 

3 

M 

126-145 

136 

I.I  0-1.  43 

1.28 

5 

F 

95-159 

120 

7.00-2.25 

2.44 

1973,  196  cm,  empty;  CAS  16302  (2383).  M,  27  May  1973.  122 
cm,  empty:  MVZ  139211  (LGB  317),  M,  24  Mar.  1970,  no  tl. 
18  otoliths.— Eumetopias:  ?(REJ  699),  F'.',  28  Sep.  1973,  192 
cm.  24  otoliths,  I  cephalopod  beak:  CAS  21400  (REJ  694),  M, 
28  Sep.  1973,  no  tl.  empty:  CAS  21398  (REJ  675),  F,  2  Aug. 
1973,  220cm,  I  otolith.— Phoca:  ''(REJ  696),  F,  28  Sep.  1973, 
133  cm,  9  otoliths.  13  cephalopod  beaks;  CAS  21424  (REJ 
689),  F,  28  Sep.  1973,  139cm,  15  Eptatretus  stoutii  eggs. 

Monterey  Co.—Zalophus:  CAS  21415  (REJ  686),  M,  22 
Sep.  1973,  no  tl,  7  otoliths;  ?(REJ  685),  M,  22  Sep.  1973,  142 
cm,  18  otoliths;  CAS  21420  (REJ  667),  M,  19  June  1973,  195 
cm,  7  otoliths;  CAS  21417  (REJ  666),  M,  19  June  1973,  256 
cm,  3  otoliths;  CAS  21416  (REJ  665),  M,  19  June  1973,  198 
cm,  48  otoliths,  40  cephalopod  beaks;  CAS  21419  (REJ  664), 
M,  19  June  1973,  151  cm,  221  otoliths:  CAS  21418  (REJ  662), 
M,  19  June  1973,  126  cm,  1  otolith,  3  cephalopod  beaks;  CAS 
21414  (REJ  647),  M,  1  Apr.  1973,  115  cm,  1  otolith,  3  cepha- 
lopod beaks;  UC  tag  6588  (REJ  244),  M,  6  June  1970,  125  cm, 
2  otoliths,  3  cephalapod  beaks;  CAS  21413  (REJ  68-40),  M,  12 
June  1968,  233  cm,*  39  otoliths.— Phoca:  CAS  21427  (REJ 
671),  M.  13  June  1973,98cm,  1  otolith;  CAS  21426  (REJ  663), 
F?.  19  June  1973,  87  cm,  empty;  CAS  21425  (REJ  646),  F,  1 
Apr.  1973,  142  cm,  10  cc  fish  bones,  3  cephalopod  beaks. — 
Callorhinus:  MVZ  153256  (REJ  645).  F,  1  Apr.  1973,  121  cm, 
empty;  ?(REJ  243),  F,  6  June  1970,  101  cm,  12  otoliths,  14 
cephalopod  beaks;  MVZ  138677  (REJ  69-36),  F,  12  July  1969, 
137  cm,  empty. 

RESULTS  • 

Sixty-eight  percent  of  the  102  stomachs  ex- 
amined in  this  study  contained  material  (Table 
1),  approximately  the  same  percentage  as  re- 
corded for  stomachs  from  collected  living  ma- 
rine mammals.  For  example,  331  of  437  stom- 
achs (76  percent)  of  the  fur  seals  taken  off 
California  in  1966  contained  food  (Marine  Mam- 
mal Biological  Laboratory  1969),  and  18  of  44 
(41  percent)  California  sea  lions  taken  recently 
in  Oregon  contained  food  (Mate  1973).  Normally 
about  60  percent  of  the  northern  sea  lions  col- 
lected during  daylight  hours  have  food  in  their 


stomachs  (Spalding  1964),  but  Mathisen  et  al. 
(1962)  found  food  in  82  percent  (114  stomachs). 
Forty  percent  of  1300  fur  seal  stomachs  exam- 
ined in  Alaska  contained  food  (Scheffer  1950a). 

Fish  otoliths  or  other  dietary  remains  were 
recovered  from  61  stomachs,  and  33  stomachs 
had  remnants  of  identifiable  invertebrates  (Ta- 
bles 3,  4,  5,  and  9).  The  items  in  the  stomachs 
included  fish  bones  and  otoliths,  parasites,  sea- 
weeds, fish  egg  cases,  cephalopod  tissue  and 
beaks,  eye  lenses  of  fish  and  cephalopods, 
rocks,  wood,  and  parts  of  other  invertebrates. 

Fitch  and  Brownell  (1968)  have  presented  a 
valid  case  for  the  use  of  otoliths  in  determination 
of  dietary  habits.  Other  investigators  also  have 
found  undigested  parts  such  as  otoliths  and 
cephalopod  beaks  in  the  forestomachs  of  ceta- 
ceans (Rae  1965;  Harrison  et  al.  1970;  Iverson 
and  Pinkas  1971;  Loeb  1972;  Smith  and  Gaskin 
1974). 

Phoca  vitulina,  Harbor  seal 

Harbor  seals  in  the  Aleutians  contained  fewer 
prey  species  than  in  other  areas  studied.  Wilke 
(1957)  examined  seven  harbor  seals  collected  in 
March  at  Amchitka  Island  and  found  that  octo- 
pus was  the  most  frequent  item,  but  that  gadid 
and  hexagrammid  fishes  made  up  the  greatest 
volume  of  food  present.  A  later  study  by  Ken- 
yon  (1965)  found  only  octopus  (Octopus  sp.)  and 
Atka-mackerel  (Pleurogrammus  monoptery- 
gius)  in  1 1  seals  at  Amchitka  Island.  He  theo- 
rized that  harbor  seals  feed  during  the  daylight 
hours  as  suggested  by  the  freshness  and  large 
volumes  of  food  in  the  stomachs.  Kenyon  failed 
to  consider  Wilke's  earlier  study  which  indicat- 


416 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


TABLE  3.     TOTAL  NUMBER  OF  FISH  OTOLITHS  FOUND  IN  22  SPECIMENS  OF  CETACEANS,  NORTH-CENTRAL  CALIFORNIA, 
1968-1973. 

Phocoena 

Phoecoenoides            Lagenorhynchus 

Delphinus 

(14) 

(2)                                    (5) 

(1) 

No. 

Percent 

No. 

Percent           No. 

Percent 

No. 

Percent 

Prey  species 

otoliths 

total 

otoliths 

total           otoliths 

total 

otoliths 

total 

Rockfishes  (Scorpaeniclae) 

1017 

(71.8) 

0 

8 

(9.2) 

0 

_ 

Northern  anchovy 

253 

(17.8) 

0 

5 

(5.7) 

0 

- 

Pacific  hake 

74 

(5.8) 

61 

(91.1)                3 

(3.5) 

0 

- 

Smelts  (Osmeridae) 

9 

(0.15) 

0 

34 

(39.0) 

1 

(9.1) 

Pacific  tomcod 

34 

(2.3) 

6 

(8.9)                0 

- 

0 

- 

Plainfin  midshipman 

-> 

(0.14) 

0 

34 

(39.0) 

0 

- 

Flatfishes  (Pleuronectidae) 

13 

(0.92) 

0 

1 

(1.2) 

0 

- 

Medusafish 

0 

- 

0 

0 

- 

10 

(90.9) 

Surfperches  (Emhiotocidae) 

9 

(0.63) 

0 

0 

- 

0 

- 

Spotted  cusk-eel 

•j 

(0.14) 

0 

0 

- 

0 

- 

Jackmackerel 

0 

- 

0 

_                   i 

(2.3) 

0 

- 

Total 

1413 

(99.68) 

67 

(100)              87 

(99.9) 

11 

(100.0) 

ed  that  the  abundant  rock  greenling  (Hexagram- 
mos  lagocephalus)  made  up  96  percent  of  food 
volume  from  harbor  seals.  Total  sample  size 
during  March  for  both  Wilke  (1957)  and  Kenyon 
(1965)  was  10  specimens,  which  is  hardly  ade- 
quate for  the  construction  of  generalities.  Yet 
Kenyon's  ( 1965)  study  is  the  basis  for  Morejohn 
and  Baltz's  (1970)  model  for  selective  feeding. 
They  looked  at  a  single  elephant  seal  and  com- 
pared its  feeding  behavior  to  that  of  harbor  seals 
at  Amchitka.  The  sample  sizes  in  these  studies 
are  barely  adequate  for  comparison,  and  are  in- 
adequate for  feeding  models. 

The  diet  of  harbor  seals  varies  greatly  with 
season  and  location  of  populations.  Pelagic,  bot- 
tom-dwelling, and  anadromous  fishes  have  all 
been  reported  in  its  diet.  Captain  Scammon 
(1874)  noted  that  the  "Leopard  Seal"  pursued 
and  devoured  small  fish. 

During  a  two-year  study  in  Alaska  (Imler  and 
Sarber  1947),  166  (41.5  percent)  of  400  harbor 
seal  stomachs  contained  identifiable  food  items. 
In  the  Copper  River  flats  of  Alaska,  67  seals  fed 
almost  entirely  on  eulachon  (Thaleichthys  pa- 
cificus).  Ninety-nine  other  specimens  from 
southeastern  Alaska  had  fed  on  walleye  pollock 
(Theragra  chalcogramma)  and  Pacific  tomcod 
(Microgadus  proximus)  (22.6  percent),  Pacific 
herring  (Clupea  harengus  pallasii)  (16.4  per- 
cent), and  flounders  (11.1  percent).  Lesser  num- 
bers (29.5  percent)  of  salmonids,  sculpins,  rock- 
fish,  blennies,  and  skates  were  reported.  Imler 


and  Sarber  (1947)  also  found  shrimp  and  octopus 
(20.6  percent)  in  harbor  seals  from  Alaska. 

Spalding  (1964)  collected  harbor  seals 
throughout  the  year  in  British  Columbia;  57  of 
the  126  stomachs  were  empty.  He  found  that 
minimal  food  was  consumed  during  the  June  to 
September  pupping  season.  In  summer,  the 
stomachs  contained  cephalopods  (35.4  percent), 
rockfish  (22.6  percent),  and  salmon  (16.1  per- 
cent). Stomachs  collected  from  September  to 
December  contained  invertebrates  (34.8  per- 
cent), herring  (10.8  percent),  and  salmon  (30.4 
percent).  Fish  of  commercial  value  composed  54 
percent  of  the  harbor  seals'  diet  on  a  yearly  ba- 
sis. 

In  Puget  Sound  also  the  harbor  seal  is  a  gen- 
eralized feeder  (Scheffer  1928).  Scheffer  and 
Sperry  (1931)  point  out  that  fishes  made  up  93.6 
percent,  molluscs  5.8  percent,  and  crustaceans 
0.6  percent  of  the  total  volume  of  harbor  seal 
stomach  contents.  Only  two  percent  of  the  har- 
bor seal  stomachs  contained  salmon  (Scheffer 
and  Slipp  1944).  Studies  in  Washington  (Scheffer 
1928;  Scheffer  and  Sperry  1931;  Scheffer  and 
Slipp  1944;  Seed  1972)  revealed  that  the  major 
prey  species  were  flatfishes:  English  sole  (Par- 
ophrys  vetulus),  flathead  sole  (Hippoglossoides 
elassodon),  Pacific  herring,  Pacific  tomcod,  Pa- 
cific hake  (Merluccius  productus),  sculpins 
(Leptocottus  armatus,  Myoxocephalus  sp.), 
walleye  pollock,  surfperches  (Cymatogaster  ag- 
gregata,  Rhacochilus  sp.),  Pacific  cod  (Gadus 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


417 


TABLE  4.     TOTAL  NUMBER  OF  FISH  OTOLITHS  FOUND  IN  38  SPECIMENS 

1968-1973. 

OF  PINNIPEDS,  NORTH-CENTRAL  CALIFORNIA, 

Phoca 

Zalophus 

Eumetopias 

Callorhinus               Mirouni><t 

(6) 

(20) 

(9) 

(1)                              (2) 

No. 

Percent 

No. 

Percent 

No. 

Percent 

No. 

Percent        No. 

Percent 

Prey  species 

otoliths 

total 

otoliths 

total 

otoliths 

total 

otoliths 

total       otoliths 

total 

Pacific  hake 

1 

(2.3) 

574 

(62.8) 

55 

(21.7) 

0 

26 

(96.3) 

Northern  anchovy 

0 

- 

218 

(23.8) 

1 

(0.4) 

12 

(100) 

- 

Rockfishes  (Scorpaenidae) 

1 

(2.3) 

57 

(6.2) 

79 

(31.2) 

0 

0 

- 

Flatfishes  (Pleuronectidae) 

4 

(9.3) 

1 

(0.1) 

44 

(17.3) 

0 

0 

- 

Spotted  cusk-eel 

0 

- 

6 

(0.6) 

30 

(11.8) 

0 

0 

- 

Plainfin  midshipman 

0 

- 

30 

(3.2) 

1 

(0.4) 

0 

0 

- 

Sahlefish 

0 

- 

0 

- 

")") 

(8.7) 

0 

0 

- 

Surfperches  (Emhiotocidae) 

18 

(41.9) 

3 

(0.3) 

0 

- 

0 

0 

- 

Pacific  herring 

0 

- 

16 

(1.7) 

1 

(0.4) 

0 

0 

- 

Lingcod 

0 

- 

1 

(0.1) 

13 

(5.1) 

0 

0 

- 

Blackhelly  eelpout 

12 

(27.9) 

0 

- 

1 

(0.4) 

0 

0 

- 

Jack  mackerel 

0 

- 

3 

(0.3) 

3 

(1.2) 

0 

0 

- 

Smelts  (Osmeridae) 

0 

- 

2 

(0.2) 

i 

(0.2) 

0 

0 

- 

Kelp  greenling 

4 

(9.3) 

0 

- 

0 

- 

0 

0 

- 

Pacific  tomcod 

2 

(4.7) 

1 

(0.1) 

0 

- 

0 

0 

- 

Brown  cat  shark 

0 

- 

0 

- 

1 

(0.4) 

0 

1 

(3.7) 

Pacific  hagfish 

1 

(2.3) 

0 

- 

0 

- 

0 

0 

- 

Chinook  salmon 

0 

- 

1 

(0.1) 

0 

- 

0 

0 

- 

Queenfish 

0 

- 

1 

(0.1) 

0 

- 

0 

0 

- 

Blacktail  snailfish 

0 

- 

o 

- 

1 

(0.4) 

0 

0 

- 

Total 

43 

(100) 

914 

(99.6) 

254 

(99.6) 

12 

(100)           27 

(100) 

nnicrocephalus),  and  lingcod  (Ophiodon  elon- 
gatus}.  These  authors  list  13  kinds  of  Crustacea 
and  4  kinds  of  mollusca  from  the  stomachs  of 
harbor  seals.  Harbor  seals  eat  flounder,  sole, 
herring,  eel,  goby,  cod,  whiting,  squid,  whelks, 
crab,  and  mussels  (King  1964).  Fishes,  squid, 
octopus,  and  shellfish  constitute  the  diet  of  har- 
bor seals  in  California  (Daugherty  1972).  Bonnot 
(1951)  indicated  that  the  fishes,  molluscs,  and 
crustaceans  consumed  by  harbor  seals  in  Cali- 
fornia are  usually  slow-moving  or  sedentary 
forms.  The  above  authors  do  not  specify  scien- 
tific names  nor  document  sources  for  their  di- 
etary information. 

I  examined  stomachs  of  12  harbor  seals  of 
which  eight  with  food  had  eaten  eight  kinds  of 
fish  (Merluccius  productus,  Microgadus  proxi- 
mus,  Lycodopsis  pacifica,  Sebastes  spp.,  Hex- 
agrammos  decagrammus,  Embiotoca  jacksoni, 
Phanerodon  furcatus,  Glyptocephalus  zachi- 
rus),  one  kind  of  octopus  (Octopus  sp.),  and  one 
kind  of  shrimp  (Crago  sp.).  Embiotocid  perch 
constituted  41.9  percent  of  my  sample,  com- 
pared with  1 1  percent  in  Washington  (Scheffer 
and  Sperry  1931).  One  harbor  seal  (REJ  689)  had 


TABLE  5.  MINIMUM  NUMBER  OF  INDIVIDUAL  FISH  AND 
CEPHALOPODS  FOUND  IN  ELEVEN  SPECIES  OF  MARINE  MAM- 
MALS, NORTH-CENTRAL  CALIFORNIA,  1968-1973. 


Cephalopods 

Fishes 

Mini- 
mum no. 
indi- 

Minimum 
no. 
indi- 

viduals 

viduals 

repre- 

No. 

repre- 

No. 

sented 

beaks 

sented 

otoliths 

Zalophus 

32 

51 

476 

922  (8*) 

Eumetopias 

11 

27 

132 

258  (5*) 

Phoca 

12 

16 

23 

43(1*) 

Callorhinus 

7 

14 

6 

12 

Mirounga 

0 

0 

13 

26 

Phocoena 

92 

168 

712 

1429(16*) 

Phocoenoides 

0 

0 

34 

67 

Lagenorhynchus 

86 

155 

45 

88(1*) 

Delphinus 

3 

4 

6 

11 

Kof>ia 

112 

217 

1 

2 

Grampus 

0 

0 

0 

0 

Totals 

355 

652 

1448 

2858(31) 

*  Unidentifiable. 

418 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


TABLE  6.  NUMBER  OF  INDIVIDUAL  FISH,  GROUPED  BY 
NICHE,  FOUND  IN  THE  STOMACHS  OF  MARINE  MAMMALS 
FROM  NORTH-CENTRAL  CALIFORNIA,  1968-1973. 

Schooling  Bottom-     Inshore 
Sample       (open      dwelling    (school- 
size         water)      (rocky)         ing) 


Zcilophus 

30 

2% 

52 

121 

Eumelopias 

19 

28 

97 

3 

Phocti 

12 

2 

24 

0 

Mirounga 

2 

13 

0 

0 

Callorhinus 

4 

0 

0 

6 

Phocoenu 

20 

52 

13 

646 

Lagenorhynchus 

7 

3 

22 

20 

Phocoenoides 

5 

33 

0 

0 

Delphinus 

1 

5 

0 

1 

Grampus 

1 

0 

0 

0 

Kogia 

1 

0 

1 

0 

ingested  Pacific  hagfish  (Eptatretus  stoutii)  egg 
cases  (15  eggs).  I  observed  that  Pacific  hagfish 
or  Pacific  lamprey  (Lampetra  tridentatd)  was  a 
prey  item  of  seals  near  river  mouths  in  northern 
California.  Scheffer  and  Sperry  (1931)  note  that 
two  percent  of  stomachs  in  which  items  oc- 
curred contained  Pacific  lamprey. 

At  birth,  harbor  seals  average  81.6  cm  in 
length  (Bigg  1969).  I  examined  two  pups  (98  and 
97  cm)  which  contained  solid  food:  an  otolith 
(Sebastes  spp.)  in  one  stomach  and  a  number  of 
shrimp  (Crago  sp.)  in  the  other.  This  suggests 
that  harbor  seals  catch  their  own  food  at  an  early 
age. 

In  summary,  knowledge  to  date  indicates  that 
harbor  seals  feed  on  shallow-water  fishes  and 
bottom-dwelling  invertebrates  (Tables  1,  6  and 
9). 

Mirounga  angustirostris,  Elephant  seal 

Pike  and  MacAskie  (1969)  reported  hagfish 
eggs  (Eptatretus)  and  probably  digested  hagfish 
remains  in  an  elephant  seal  from  Canada.  Huey 
(1930)  pointed  out  that  several  of  the  fish  species 
found  in  elephant  seal  stomachs  inhabit  water 
from  50  to  120  fathoms  (ca.  91  to  219  m)  deep. 
Small  sharks  (Squalus,  Cephaloscyllium),  skates 
(Raja),  rays  (Myliobatidae),  and  ratfish  (Hydro- 
lagus)  have  been  reported  as  food  items  in  ele- 
phant seals  (Kenyon  and  Scheffer  1955;  King 
1964;  Daugherty  1972;  Seed  1972).  Huey  (1925) 
found  that  three  of  four  Guadalupe  Island  ele- 
phant seals  contained  squid  (Loligo). 

Contrary  to  the  report  of  Morejohn  and  Baltz 
(1970),  teleost  fishes  had  been  reported  from  el- 


ephant seals.  Huey  (1930)  reported  a  single  bass, 
and  Daughterly  (1972)  stated  that  elephant  seals 
eat  rockfish.  Freiburg  and  Dumas  (1954)  found 
a  dead  adult  elephant  seal  in  Oregon  which  may 
have  died  from  bones  of  Pacific  hake  (Merluc- 
cius  productus)  blocking  the  internal  nasal  re- 
gion. 

More  information  is  needed  on  fish  ecology 
and  fish  population  abundance  before  the  nature 
of  feeding  behavior  of  elephant  seals  can  be  clar- 
ified. The  elephant  seal  may  be  a  selective  feeder 
(Morejohn  and  Baltz  1970),  but  seasonal,  sex, 
and  age  factors  need  to  be  resolved.  Commercial 
catches  are  the  reference  for  fish  abundance  by 
Morejohn  and  Baltz.  However,  commercial  fish 
catches  are  a  poor  indication  of  fish  abundance 
when  most  reported  fish  are  of  commercial  value 
(Bell  1971).  Anyone  accompanying  commercial 
vessels  realizes  that  many  unreported  fish  are 
caught  besides  those  marketed.  All  fishes  More- 
john and  Baltz  found  were  bottom  or  rock- 
dwelling  species  (Chilara,  Porichthys,  Sebastes, 
and  Lyopsetta  or  Glyptocephalus). 

Elephant  seals  may  feed  during  particular  pe- 
riods of  the  day.  If  they  are  nocturnal  feeders, 
they  may  catch  hake  high  in  the  water  column, 
but  diurnal  feeding  would  suggest  that  hake  are 
taken  in  deep  water  (Nelson  and  Larkins  1970). 
Studies  are  needed  to  determine  when  and 
where  elephant  seals  feed.  Large  fish  otoliths 
may  indicate  offshore  feeding  by  the  elephant 
seal. 

To  date,  few  elephant  seals  have  been  exam- 
ined for  stomach  contents.  One  of  my  seals,  an 
immature  female  (HSC-1970)  found  dead  in 
Humboldt  County  on  5  May  1970,  contained  26 
otoliths  from  Pacific  hake,  representing  at  least 
12  adult  fish.  Hake  migrate  north  along  the  coast 
in  spring  and  summer  and  might  be  expected  to 
be  present  in  marine  mammal  diets  at  that  time 
of  year.  Pacific  hake  form  massive  schools  just 
above  the  bottom  and  show  a  pronounced  daily 
vertical  migration  (Nelson  and  Larkins  1970). 
Most  adult  hake  are  located  beyond  the  conti- 
nental shelf  at  depths  of  230-410  m.  Although 
Pacific  hake  are  present  in  commercially  abun- 
dant numbers,  they  are  not  being  exploited  cur- 
rently by  American  fisheries.  Pacific  hake 
ranked  second  in  abundance  in  California  larval 
surveys  (Ahlstrom  1965). 

My  other  specimen,  also  an  immature  female, 
was  found  in  Trinidad,  Humboldt  County,  19 
Feb.  1969  (HSC-69-6).  It  contained  a  single  egg 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


419 


case  of  the  brown  cat  shark  (Apristurus  brun- 
neus),  which  was  identified  using  the  description 
and  photos  in  Cox  (1963).  This  species  of  shark 
is  found  in  deep  water  from  British  Columbia  to 
Baja  California. 

Eumetopias  jubatus,  Northern  sea  lion 

The  northern  sea  lion  has  been  studied  more 
intensively  than  most  other  North  Pacific  pin- 
nipeds. This  sea  lion  occurs  along  the  eastern 
Pacific  coasts  from  Alaska  to  California,  where 
its  relationships  to  commercial  fisheries  have 
been  studied  extensively.  In  general,  these  stud- 
ies reveal  that  fish  and  cephalopods  are  the  pre- 
ferred foods. 

In  1899  L.  L.  Dyche  inspected  25  sea  lion 
stomachs  from  near  Monterey  Bay,  California. 
All  specimens  referred  to  by  Dyche  contained 
squid  or  octopus. 

The  original  manuscript  Dyche  sent  to  C.  H. 
Merriam  in  1901  did  not  specify  which  sea  lion 
was  involved.  This  partially  handwritten  docu- 
ment mentioned  only  the  common  identification 
"California  Sea  Lion"  (Unpublished  manu- 
script, Dyche  1901.  C.  H.  Merriam  file  at  Mu- 
seum of  Vertebrate  Zoology,  University  of  Cal- 
ifornia, Berkeley).  When  Merriam  (190 la,  1901b) 
first  published  Dyche's  data,  he  did  not  indicate 
whether  the  northern  sea  lion  (Eumetopias)  or 
the  California  sea  lion  (Zalophus)  was  involved. 
Later,  Dyche  (1903)  used  the  name  "Zalophus 
californianus  Lesson"  beneath  the  general  title 
of  "Food  for  California  Sea  Lions,"  suggesting 
that  California  sea  lions  were  examined.  Thus 
it  was  long  believed  that  all  sea  lions  examined 
by  Dyche  were  Zalophus.  Briggs  and  Davis 
(1972)  have  pointed  out  that  at  least  some  of 
Dyche's  specimens  were  Eumetopias  jubatus. 
Since  only  7  of  these  25  sea  lions  deposited  at 
the  University  of  Kansas  are  extant  today,  it  is 
not  possible  to  make  positive  identifications  for 
all  of  Dyche's  specimens  (R.  S.  Hoffmann,  Cu- 
rator Univ.  of  Kansas,  pers.  comm.  [1973]).  Al- 
though the  7  extant  specimens  are  northern  sea 
lions,  it  is  possible  that  all  25  of  the  originally 
collected  specimens  may  not  have  been  this 
species. 

Northern  sea  lions  have  been  reported  to  feed 
at  night  (Rowley  1929;  Bonnot  1951;  Mathisen 
1959;  Mathisen  et  al.  1962;  Spalding  1964;  Seed 
1972;  Mate  1973).  Fiscus  and  Baines  (1966) 
sighted  feeding  groups  of  up  to  several  thousand 
animals  8-22  km  out  in  Unimak  Pass,  Alaska. 


It  would  be  of  interest  to  know  if  the  individuals 
observed  feeding  returned  to  the  hauling-out 
area  each  afternoon  with  engorged  stomachs. 
Daytime  feeding  behavior  has  been  noted  by 
many  fishermen,  specifically  long-line  and  drag- 
boat  operators  (Kenyon  1952a). 

An  interesting  examination  at  the  cannery 
dock  at  the  mouth  of  the  Klamath  River  was 
reported  by  Bonnot  (1951).  Two  half-grown 
northern  sea  lions  were  killed,  and  only  lam- 
preys were  found  in  their  stomachs.  J.  C.  Snyder 
also  examined  sea  lions  at  river  mouths  and 
identified  the  remains  of  lampreys  in  their  stom- 
achs (Kenyon  1952b).  More  recently,  Jameson 
and  Kenyon  (1978)  reported  that  82  percent  of 
observed  feeding  behavior  at  the  Rogue  River, 
Oregon,  was  on  lampreys. 

Rutter  et  al.  (1904)  presented  data  on  18  north- 
ern sea  lions  (6  males  and  12  females)  from 
north-central  California.  The  eight  female  sea 
lions  containing  food  were  taken  from  Afio  Nue- 
vo  Island  in  July  or  August.  Five  male  sea  lions 
from  Pt.  Arena  ate  at  least  147  fish  but  only  5 
squid.  The  majority  of  their  food  was  fish  (257 
fish  present  in  the  13  animals).  The  only  ceph- 
alopod  material  was  from  five  sea  lions  from 
both  areas.  This  low  incidence  of  cephalopods 
does  not  agree  with  the  findings  of  Dyche  (1903), 
but  the  identity  of  Dyche's  specimens  is  in 
doubt.  In  contrast,  in  southern  California  Rutter 
et  al.  (1904)  found  only  39  fish  but  thousands  of 
cephalopods  in  a  sample  of  24  Zalophus.  They 
concluded  that  the  northern  sea  lion  feeds  chief- 
ly on  fish,  and  the  California  sea  lion  on  ceph- 
alopods, and  that  both  feed  opportunistically. 

Bonnot  (1928)  examined  two  northern  sea  lion 
stomachs  from  San  Miguel  Island,  California,  on 
20  June  1928.  The  adult  male  was  empty  but  the 
female  contained  three  greenish  eggs  of  a  skate 
or  shark.  Bonnot  stated  that  this  was  an  old, 
blind  sea  lion  and  perhaps  she  was  dying.  In 
northern  California,  at  the  Saint  George  Reef 
rookery  (Del  Norte  County),  Bonnot  found 
thousands  of  shells  of  a  tiny  pelecypod  embed- 
ded in  sea  lion  feces.  Other  investigators  have 
not  reported  analysis  of  scats,  perhaps  because 
of  the  difficulty  of  locating  adequate  samples 
(Bonnot  1928). 

The  literature  contains  a  composite  list  of  32 
fishes  from  stomachs  of  northern  sea  lions  (Ta- 
ble 7).  Because  of  the  nonspecific  categories  of 
some  fish  names,  I  have  not  attempted  to  pro- 
duce an  accurate  species  list.  Only  the  papers 


420 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


TABLE  7.     SUMMARY  OF  FOOD  CONTENTS  OF  Eumetopias  (modified  from  Pike  (1958)  with  recent  additions). 


Season 
Locality                                          and  year                    Source 

No.  of  stomachs 
(size  and/or  sex) 

No.  of  stomachs, 
kinds  of  food 

St.  Paul  I.,  Alaska                    July  1949-1951                             (6) 

~>~> 
(2  females) 

19  —  empty,  1  —  ceph. 
beak,  1-10  Ibs., 
sandlance  trace-starry 
flounder,  sculpin,  1-20 
Ibs.  halibut,  cod, 
flounder,  pollack, 
4  —  large  stones 

Sitka  to  Kodiak  I.,                   May,  July,  Aug.                         (4) 
Alaska                                        1945-1946 

23 
(adults) 

8-empty,  15  —  salmon, 
cod,  halibut,  pollack, 
4  —  octopus 

Kodiak  to  Krenitzin                 May-July  1959                           (8) 
I.,  Alaska 

382 
(372  males, 
10  females) 

326—  empty.  28—  fish 
(7spp.),  21— 
invertebrates,  20  — 
squid  and  octopus, 
10—  no  ID  fish,  154— 
stones  and  gravel 

Chernaburg  I.,                          May,  July  1958                          (7) 
Alaska 

114 
(46  males, 
51  females, 
17  yearlings) 

20—  empty,  31—  fish 
(7  spp.),  61— 
invertebrates,  73  —  rock 
and  sand,  24  —  no  ID 
fish,  5  —  milk 

Beresford  I.,  B.C.                    Aug.  1913                                   (2) 

3 
(adults) 

2  —  empty,  1  —  salmon, 
"cod"  and  "bass" 

Scott  I.,  B.C.                            June,  July  1956                          (5) 

56 
(adults) 

50  —  empty,  4  —  fish  and 
squid,  1  —  herring, 
1  —  octopus 

Scott  I.,  B.C.                            June  1957                                    (5) 

8 
(4  females, 
4  males) 

6  —  empty,  1  —  salmon, 
1—  no  ID  fish 

Barkley  Sound,  B.C.                Dec.  1915                                    (2) 

14 
(1  1  males, 
3  females) 

12  —  herring,  1  —  clam- 
shell, 1  —  crab,  octopus 

Barkley  Sound,  B.C.                Feb.,  Apr.  1958                         (5) 

14 
(13  females, 
1  male) 

6  —  empty,  3  —  herring, 
2—  rockfish.  2—  fish/ 
octopus,  1  —  octopus, 
skate,  hake 

Isnor  Rock,  B.C.                      July,  Aug.  1957                          (5) 

3 
(young  males; 

1  —  rockfish,  1  —  squid  and 
rockfish.  1  —  squid 

British  Columbia                      Feb.  -Dec.  1959                          (9) 

393 
(equal  numbers, 
males/females) 

213—  empty,  75—  fish  (17 
spp.),  49—  no  ID,  milk 
or  kelp 

Offshore  California-                Mar.,  Sep.  1958-1  963              (10) 
N.  Pacific 

34 
(7  males, 
15  females) 

1  1  —  lost  at  sea,  22  —  fish 
(15  spp.).  2—  no  ID 
fish,  1  —  clamshell  and 
fish,  9  —  rocks  and 
pebbles 

Pt.  Arena  and                           July-Aug.  1901                           (1) 
Afio  Nuevo  I.,  Calif. 

18 
(6  males, 
12  females) 

5  —  empty,  13  —  -fish,  6  — 
squid  and  octopus 

JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


421 


TABLE  7.     CONTINUED. 


Locality 

Season 
and  year 

Source 

No.  of  stomachs 
(size  and/or  sex) 

No.  of  stomachs, 
kinds  of  food 

Afio  Nuevo  I.,  Calif. 


"several  years  ago' 
prior  to  1918 


(3) 


15 

(14  females, 
1  young  male) 


7 — empty,  8 — rock, 
sardines,  salmon, 
3 — fish  and  squid 


North-Central 
California 


1968-1973 


(11) 


19 

(7  males, 
12  females) 


10— empty,  9— fish,  7— 
squid  and  octopus,  2— 
rocks 


(1)  Rutteret  al.  1904. 

(2)  Newcombeet  al.  1918. 

(3)  Starks  1918. 

(4)  Imler  and  Sarber  1947. 

(5)  Pike  1958. 

(6)  Wilke  and  Kenyon  1952. 

(7)  Mathisen  et  al.  1962. 

(8)  Thorsteinson  and  Lensink  1962. 

(9)  Spalding  1964. 

(10)  Fiscus  and  Baines  1966. 

(11)  Current  study  1973. 


of  Wilke  and  Kenyon  (1952),  Spalding  (1964), 
and  Fiscus  and  Baines  (1966)  present  scientific 
names  and  volumetric  determinations  which  en- 
able me  to  present  a  well-documented  dietary 
list.  Pike  (1958)  also  presented  a  table  with  stom- 
ach contents.  All  of  these  data  are  updated  and 
presented  as  Table  7. 

My  study  adds  10  genera  of  fishes  to  those 
previously  reported  from  northern  sea  lions,  as 
follows:  Microstomus,  Parophrys,  Careproctus, 
Lyopsetta,  Eopsetta,  Glyptocephalus,  Por- 
ichthys,  Engraulis,  Spirinchus,  and  Chilara. 
These  10  genera  constituted  31.1  percent  of  the 
otoliths  found  in  Eumetopias. 

All  northern  sea  lion  stomachs  which  con- 
tained cephalopod  beaks  also  had  remains  of 
from  2  to  13  species  of  fish  (Table  5).  Four 
species  of  cephalopods  were  identified:  Loligo 
opalescens,  Octopus  sp.,  Chiroteuthis  sp.,  and 
Onychoteuthis  sp.  Most  beaks  were  so  thor- 
oughly digested  that  specific  identifications  were 
impossible.  One  male  sea  lion  (REJ  629)  had  eat- 
en at  least  3  octopus  plus  13  species  of  fish. 
More  surprising  than  the  variety  of  prey  eaten 
was  the  fact  that  the  sea  lion  had  been  eating 
during  the  breeding  season  when  most  Eume- 
topias males  fast  (Spalding  1964). 

In  my  study  demersal  fish  were  found  in  six 
of  the  nine  stomachs  containing  fish.  When  the 
127  identified  fishes  from  northern  sea  lions  are 


grouped  according  to  schooling  (open- water), 
bottom-dwelling  (rocky),  and  inshore-schooling 
species  (Table  6),  it  is  apparent  that  the  northern 
sea  lion  feeds  mainly  on  bottom-dwelling  fishes. 
The  rather  high  incidence  of  rocks  in  the  stom- 
achs also  suggests  a  bottom-feeding  habit  (Ta- 
bles 1  and  9). 

Zalophus  calif ornianus,  California  sea  lion 

California  sea  lions  make  annual  north-south 
migrations  along  the  Pacific  Coast  of  North 
America.  Adult  and  subadult  males  move  north- 
ward during  September  and  October  after  the 
breeding  season  (Bonnot  1928;  Fry  1939;  Orr 
and  Poulter  1965,  1967;  Bartholomew  1967;  Pe- 
terson and  Bartholomew  1967;  Peterson  and 
LeBoeuf  1969;  Odell  1971)  and  return  south  in 
March  to  the  more  southern  breeding  rookeries. 

Virtually  nothing  is  known  about  the  feeding 
behavior  of  migrating  California  sea  lions.  Pe- 
terson and  LeBoeuf  (1969)  indicated  that  influx- 
es of  sea  lions  into  northern  areas  are  correlated 
with  periods  of  abundance  of  food,  but  they  did 
not  document  their  statement. 

No  published  studies  have  been  reported  on 
California  sea  lions  between  Monterey  Bay  and 
the  Oregon  border.  B.  R.  Mate  (Oregon  State 
Univ.,  pers.  comm.  [1971])  collected  44  male 
California  sea  lions  in  Oregon,  but  he  has  not 
yet  identified  the  fish  otoliths.  From  the  data 


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presented  by  Mate  (1973),  one  can  calculate  that 
40.9  percent  of  the  combined  sample  of  delib- 
erately collected  sea  lions  contained  food.  My 
method  of  obtaining  stomach  data  showed  that 
66.6  percent  of  the  beached  California  sea  lions 
contained  food. 

Fiscus  and  Baines  (1966)  examined  six  Cali- 
fornia sea  lions  taken  during  a  recent  study  on 
fur  seals.  The  stomach  volume  was  recorded; 
thus  the  estimate  for  feeding  rates  and  food  in- 
take to  body  weight  can  be  determined  more 
clearly.  Loligo  opalescens  (common  squid),  un- 
identified squid,  northern  anchovy,  and  Pacific 
hake  were  found  in  these  pelagic  California  sea 
lions. 

Several  observations  have  been  made  of  feed- 
ing sea  lions.  Fink  (1959)  reported  a  single  ob- 
servation, on  25  Feb.  1959,  in  Monterey  Bay  of 
California  sea  lions  attacking  a  school  of  Pacific 
sardines.  Although  feeding  on  sardines  is  surely 
not  uncommon,  his  vivid  description  of  harbor 
porpoises  controlling  the  fish  school  and  the  sea 
lions  feeding  on  the  periphery  of  the  school  is 
the  only  published  account  of  such  behavior. 
Ryder  (1957)  reported  feeding  aggregations  of 
pinnipeds  and  birds.  At  the  Farallon  Islands, 
California,  an  adult  male  California  sea  lion  was 
seen  repeatedly  eating  several  jackmackerel 
during  the  daylight  hours  on  17  and  19  Septem- 
ber 1973  (T.  James  Lewis  and  Barbara  Lewis, 
Point  Reyes  Bird  Observatory,  pers.  comm.).  At 
Cerros  (Cedros)  Island,  Baja  California,  a  fe- 
male California  sea  lion  was  observed  feeding 
beneath  the  surface  by  Bonnot  (1932b),  who 
watched  this  cow  eat  at  least  six  large  flying  fish 
(Exocoetidae). 

Scheffer  and  Neff  (1948)  noted  that  the  anal- 
ysis of  only  58  California  sea  lion  stomachs  had 
been  reported  in  the  literature.  Of  these, 
Dyche's  25  specimens  from  the  Monterey  area 
are  either  misidentified  or  of  questionable  iden- 
tity. Scheffer  and  Neff  examined  four  sea  lion 
stomachs  from  southern  California.  Two  fe- 
males were  empty,  but  the  other  female  con- 
tained evidence  of  at  least  21  small  squid.  The 
single  male  found  dead  near  La  Jolla  on  26  Nov. 
1943  had  36  nearly  whole  Pacific  herring  plus 
fragments  representing  30  other  herring.  All  Cal- 
ifornia sea  lions  analyzed  by  Scheffer  and  Neff 
were  from  south  of  Point  Conception. 

Rutter  et  al.  (1904)  examined  stomachs  of  24 
California  sea  lions,  13  with  food,  in  July  and 


August  1901  at  southern  California  localities. 
The  eight  females  each  had  100  to  300  small 
squid  parts.  Squid  pens  were  food  remnants  in 
the  stomachs  of  three  of  the  five  males.  The  re- 
maining stomach  contents  consisted  of  hake, 
rockfish,  ratfish,  unidentified  small  fish,  and 
milk.  Bonnot  (1928)  reported  on  these  same 
specimens  and  stated  that  5  had  eaten  fish  and 
11  had  eaten  squid.  Starks  (1918)  pointed  out 
that  stomachs  of  two  of  the  breeding  bulls  that 
Rutter  et  al.  examined  did  not  contain  any  food. 

According  to  a  recent  survey  by  scientists  at 
the  Scripps  Institution  of  Oceanography  and 
presented  to  the  California  Senate  Fact  Finding 
Committee  on  Natural  Resources  (Anderson 
1960),  24  of  30  stomachs  from  Zalophus  con- 
tained food.  The  only  identified  items  were  fish 
otoliths,  but  many  unidentified  cephalopod 
beaks  also  were  present.  Carl  L.  Hubbs  (Scripps 
Institution  of  Oceanography)  and  John  E.  Fitch 
(California  Dept.  of  Fish  and  Game,  Long 
Beach,  pers.  comm.  [1973])  revealed  that  this 
study  was  done  by  the  late  Art  Kelly  in  southern 
California  and  northern  Mexico.  The  424  fish 
otoliths  which  Kelly  recovered  were  identified 
by  Fitch  as  representing  24  kinds  of  fishes.  Pa- 
cific hake  were  found  in  17  of  the  24  stomachs 
and  constituted  48. 1  percent  of  the  total  otoliths. 
The  other  fishes  were  cusk-eels,  midshipmen, 
and  species  of  rockfish.  Sixty-six  (15.6  percent) 
of  the  otoliths  were  of  Pacific  mackerel,  ancho- 
vy, perch,  and  white  croaker.  These  sea  lions 
also  fed  extensively  on  squid  and  octopus,  as 
represented  by  beaks  in  their  stomachs. 

Briggs  and  Davis  (1972)  spent  500  hours 
aboard  sport  and  commercial  salmon  boats  in 
Monterey  Bay  from  14  April  to  22  September 
1969.  They  observed  seven  instances  of  preda- 
tion  on  salmon  by  California  sea  lions.  Of  the 
hooked  fish,  4.1  percent  were  lost  to  sea  lions. 
I  found  10  dead  California  sea  lions  along  Mon- 
terey Bay  during  the  commercial  salmon  season 
(April  to  September).  I  also  located  an  additional 
20  California  sea  lion  carcasses  farther  north 
(Figure  1).  I  found  a  single  female  California  sea 
lion  north  of  Monterey  Bay.  Only  one  salmon 
otolith  was  present  among  the  922  otoliths  (461 
fish  represented)  from  my  sample  of  California 
sea  lions. 

Male  California  sea  lions  feed  on  a  variety  of 
schooling  fishes  (Pacific  hake,  anchovy,  rock- 
fish,  flatfish,  cusk-eel,  midshipmen,  herring, 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


423 


lingcod,  jackmackerel,  salmon,  and  osmerids). 
Pacific  hake  and  anchovy  make  up  86.6  percent 
by  frequency  of  occurrence  of  otoliths  from  sea 
lion  stomachs.  Schooling  fishes,  both  inshore 
and  open- water  types  (417  otoliths)  were  found 
in  California  sea  lions  (Table  6). 

Callorhinus  ursinus,  Northern  fur  seal 

Wilke  and  Kenyon  (1957)  identified  five 
species  of  fish  from  204  seals  (114  with  food) 
collected  from  the  Bering  Sea  and  St.  Paul  Is- 
land. Seals  collected  at  sea  contained  large  num- 
bers of  capelin  (Mallotus  villosus)  and  walleye 
pollock  (Theragra  chalcogrammd) .  Only  one 
salmon  (Oncorhynchus  sp.)  was  found  in  the 
stomachs.  Three  seals  killed  on  land  at  St.  Paul 
Island  collectively  held  one  salmon  (Oncorhyn- 
chus sp.),  one  walleye  pollock,  and  two  sandfish 
(Trichodon).  Indian  hunters  took  41  seals  (13 
empty)  30  miles  (about  48  km)  off  Washington 
in  1930.  Although  the  stomach  contents  were 
digested,  squid  eyes  and  beaks  were  reported 
from  21,  and  identifiable  herring  vertebrae  were 
present  in  15  stomachs  (Schultz  and  Rafn  1936). 
Clemens  and  Wilby  ( 1933)  looked  at  25  stomachs 
from  the  west  coast  of  Vancouver  Island  and 
reported  that  8  contained  squids,  9  had  sal- 
mon, and  the  rest  had  small  schooling  fish. 
No  data  on  the  volume  of  the  stomachs  or  the 
number  of  empty  stomachs  were  reported. 

Hanna  (1951)  recorded  fur  seals  in  the  Gulf  of 
the  Farallons  during  February  and  April.  Seals 
were  described  as  competitors  with  W.  I.  Follett 
as  he  dip-netted  for  myctophid  fish  (Tarleton- 
beania,  Symbolophorus),  sablefish  (Anoplo- 
pomafimbria),  and  red  Irish  lord  (Hemilepido- 
tus  hemilepidotus).  Hanna  also  theorized  that 
fur  seals,  porpoises,  and  sea  birds  were  all  feed- 
ing on  pteropods  (Mollusca). 

More  recently  437  stomachs  from  pelagic  fur 
seals  taken  off  California  were  examined  (Ma- 
rine Mammal  Biological  Laboratory  1969).  An- 
chovy, saury,  hake,  and  squid  constituted  98 
percent  of  the  total  food  volume. 

Scheffer  (1950a)  reviewed  the  dietary  litera- 
ture on  fur  seals.  He  presented  data  on  only  two 
northern  fur  seal  stomachs  from  California,  one 
of  which  contained  an  unidentified  bird  and  the 
other  had  fed  on  Pacific  saury  (Cololahis  saira). 

Fur  seals  found  on  California  beaches  usually 
have  little  food  in  their  stomachs.  Likewise  a  fur 
seal  from  Southeast  Farallon  Island  contained 


no  food  in  its  stomach  (REJ  212),  but  had  seven 
isopods  (Riggia?)  which  are  external  parasites 
on  fish.  One  beach-cast  fur  seal  had  7  beaks  of 
the  common  squid  and  12  northern  anchovy 
otoliths  in  its  stomach  (REJ  243). 

Phocoena  phocoena,  Harbor  porpoise 

Although  the  diet  of  harbor  porpoises  in  Cal- 
ifornia waters  is  poorly  known,  herring,  small 
cods,  soles,  and  squid  are  food  items  of  harbor 
porpoises  generally  (Ridgway  1972).  Scammon 
(1874)  wrote,  "They  feed  upon  fish,  and  are  oc- 
casionally taken  in  seines  that  are  hauled  along 
the  shores  of  San  Francisco  Bay  by  the  Italian 
fishermen." 

Harbor  porpoises  seldom  are  sighted  more 
than  20  miles  (about  32  km)  offshore  and  usually 
are  seen  near  harbor  entrances  (Fiscus  and  Nig- 
gol  1965).  Local  fish  abundance  and  seasonal 
fish  movements  affect  the  diet  of  these  porpoises 
(Rae  1965). 

In  Scottish  waters,  Rae  (1965)  examined  45 
porpoises  from  November  to  March  and  7  ad- 
ditional ones  in  the  summer  months.  A  few  in- 
vertebrates were  found  which  Rae  thought  might 
have  been  taken  incidentally  with  other  food 
items.  Fish  or  fish  remains  were  recognized  in 
41  of  the  43  stomachs  with  recognizable  food. 
One  of  the  two  remaining  stomachs  contained 
milk  and  the  other  the  remains  of  a  very  small 
cephalopod.  Ten  species  of  fish  were  present, 
with  herring  (Clupea  harengus)  and  whiting 
(Gadus  merlangus)  the  most  common.  Most  fish 
were  less  than  25  cm  in  length,  with  the  largest 
individuals  35  cm.  These  harbor  porpoises  had 
been  trapped  in  nets  set  for  cod  or  salmon. 

British  naturalists  have  recorded  food  habits 
of  harbor  porpoises  in  the  North  Sea  for  more 
than  100  years,  and  small  fish,  mainly  clupeoids 
(65  percent)  and  gadoids  (30  percent),  consti- 
tute the  major  foods  taken  (Rae  1965).  Rae  con- 
cluded that  harbor  porpoises  take  pelagic  forms 
of  fish.  Tomilin  (1957)  found  benthic  fish  pre- 
dominating in  the  diet  of  harbor  porpoises  from 
the  Black  Sea  and  Sea  of  Azov.  In  a  study  in 
the  Bay  of  Fundy,  small  schooling  fishes  (Clu- 
pea harengus,  Gadus  morhua,  Scomber  scom- 
brus)  were  principal  food  items  for  harbor  por- 
poises (Smith  and  Gaskin  1974).  These  schooling 
fishes  accounted  for  78  percent  of  the  total  diet. 

Tomilin  (1957)  listed  dietary  items  of  harbor 
porpoises  in  the  Black  Sea,  where  4000  stom- 


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achs  were  inspected.  These  porpoises  fed  upon 
eight  benthic  species  and  six  pelagic  species  of 
fish.  The  pelagic  fish  were  consumed  when  they 
occurred  in  large  dense  schools.  One  of  two  har- 
bor porpoises  found  at  Pt.  Barrow,  Alaska,  con- 
tained bones  of  whitefish  (Leucichthys)  (Hall 
and  Bee  1954).  Pike  and  MacAskie  (1969)  re- 
ported a  Canadian  harbor  porpoise  caught  in  a 
gill  net.  This  animal  had  one  herring  in  its  stom- 
ach. 

Scheffer  (1953)  inspected  a  female  porpoise 
from  Grays  Harbor  County,  Washington,  which 
had  fed  on  37  capelin.  Its  death  apparently  was 
caused  by  an  American  shad  blocking  its  throat 
passages.  Five  years  earlier  Scheffer  had  found 
on  the  same  beach  another  porpoise  which  also 
had  choked  to  death  on  a  fish.  A  female  porpoise 
from  Port  Townsend,  Washington,  May  1950, 
had  eaten  five  Pacific  herring  (Wilke  and  Ken- 
yon  1952). 

Orr  (1937)  reported  that  a  porpoise  apparently 
choked  to  death  on  a  gray  smoothhound  shark 
(Mustelus  californicus).  This  single  report  of 
sharks  as  a  food  item  may  be  abnormal.  Fink 
(1959)  observed  for  30  minutes  several  hundred 
harbor  porpoises  feeding  on  a  school  of  Pacific 
sardines  northwest  of  Pt.  Pinos,  Monterey  Bay. 
He  vividly  described  the  attack  and  herding  of 
the  sardines,  and  counted  from  5  to  12  fish  eaten 
by  an  individual  porpoise  in  its  attack  through 
the  sardines. 

In  my  study,  juvenile  rockfish  constituted  71.8 
percent  of  the  diet  of  harbor  porpoises  (Table 
3).  Five  porpoises  collected  in  June  and  July 
contained  1017  rockfish  otoliths.  Northern  an- 
chovy was  the  second  most  frequent  fish  and 
was  found  in  seven  porpoises  from  April  to  Au- 
gust. Over  two-thirds  of  all  fish  found  in  stom- 
achs of  harbor  porpoises  live  in  open  water  or 
are  inshore  schooling  species  (Table  6).  Juvenile 
Pacific  hake,  Pacific  tomcod,  rockfish,  and 
northern  anchovy  accounted  for  97  percent  of 
all  stomach  otoliths  found  during  my  investiga- 
tion (Table  3). 

Invertebrate  remains  were  found  in  8  of  the 
20  porpoises  examined  (Table  9).  A  total  of  141 
Loligo  opalescens  beaks  was  identified  from 
168  cephalopod  fragments.  These  beaks  repre- 
sent at  least  92  individual  cephalopods  com- 
pared to  at  least  712  fish  represented  by  1429 
otoliths  (Table  5).  One  harbor  porpoise  (CAS 
2392)  had  13  intact  Loligo  and  an  additional  35 
pairs  of  beaks  in  its  stomach.  The  intact  bodies 


of  the  13  squids  indicated  harbor  porpoises  do 
not  chew  this  food  item. 


Phocoenoides  dalli,  Dall's  porpoise 

This  porpoise  is  much  more  common  than  ear- 
ly records  indicate.  Brownell  (1964)  reported  its 
occurrence  in  southern  California  waters  in  the 
winter.  Lustig  (1948)  saw  10  or  12  porpoises 
feeding  on  baitfish,  anchovy  or  sauries,  on  13 
July  1939,  in  the  Anacapa  Passage. 

Deep-water  benthic  fish  and  bathypelagic  ceph- 
alopods were  reported  as  major  food  items  from 
a  large  sample  of  Dall's  porpoise  stomachs  from 
Japan  (Wilke  and  Nicholson  1958).  Eleven  per- 
cent of  the  food  volume  was  squid:  Watasenia, 
Oinmastrephes,  and  unidentified  genera.  Myc- 
tophidae  (lanternfishes)  composed  70  and  73 
percent  of  the  stomach  contents  in  1949  and 
1952,  respectively. 

Cowan  (1944)  took  five  (3  males  and  2  fe- 
males) Dall's  porpoise  off  the  coast  of  British 
Columbia  in  the  summer  of  1939.  Four  of  the 
five  stomachs  were  full  of  herring.  Pike  and 
MacAskie  (1969)  examined  three  males  and  two 
females  from  British  Columbia,  and  they  too 
found  mostly  herring  or  squid  in  three  stomachs. 

Scheffer  (1953)  recorded  the  stomach  contents 
of  six  Dall's  porpoises  from  Monterey  and 
northward.  These  contained  Pacific  hake,  squids 
(Loligo  opalescens  and  unidentified  species), 
jackmackerel,  and  unidentified  fish.  Two  Dall's 
porpoises  from  Alaska  had  fed  only  on  capelin. 
Brown  and  Norris  (1956)  mentioned  anchovy  as 
a  food  item  of  the  Dall's  porpoise.  An  adult  por- 
poise taken  in  southern  California  waters  had 
eaten  at  least  14  Pacific  hake,  2  jackmackerel, 
and  13  cephalopods  (Norris  and  Prescott  1961). 
These  authors  also  noted  the  porpoise  circling 
amid  schools  of  sauries,  probably  feeding.  Fis- 
cus  and  Niggol  (1965)  observed  Dall's  porpoises 
off  the  north  coast  and  collected  five  specimens 
off  Cape  Mendocino,  California.  Three  females 
and  one  of  the  males  had  only  squid  beaks  in 
their  stomachs;  the  stomach  of  the  other  male 
was  empty. 

The  Dall's  porpoise  is  present  in  Monterey 
Bay  all  year.  Stomach  samples,  examined  each 
month,  indicated  that  Pacific  hake,  rockfish,  and 
squid  are  important  food  items.  Loeb  (1972)  ex- 
amined 25  stomachs  of  Dall's  porpoise  from 
Monterey  Bay  and  found  Pacific  hake  in  23, 
squid  in  16.  Most  of  the  cephalopods  present 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


425 


were  Loligo,  with  lesser  numbers  of  Abraliop- 
sis,  Gonatus,  Onychoteuthis,  and  Octopus.  Pa- 
cific hake,  juvenile  rockfish,  and  squid  made  up 
93  percent  of  the  total  diet  of  the  Dall's  porpoise 
from  Monterey  Bay  (Table  5  in  Loeb  1972). 

On  28  June  1973,  an  immature  female  Dall's 
porpoise  was  found  on  the  beach  north  of  the 
University  of  California  Marine  Station  at  Bo- 
dega Bay.  Presumably  this  animal  (REJ  670,  102 
cm  total  length)  was  dependent  on  its  mother  for 
nourishment,  although  no  milk  was  noted  in  its 
stomach. 

I  examined  four  adult  Dall's  porpoises  (2 
males,  2  females),  and  only  two  had  identifiable 
food  remains.  One  (REJ  674)  had  61  Pacific  hake 
otoliths  representing  31  fish  in  its  stomach,  and 
the  other  (CAS  2335)  contained  6  juvenile  Pa- 
cific tomcod.  The  two  other  porpoises  had  emp- 
ty stomachs  (CAS  2384,  REJ  678). 

Loeb  (1972)  did  not  mention  Pacific  tomcod 
as  a  dietary  item  from  Dall's  porpoises  but  did 
note  a  wide  variety  of  fishes  (15  species)  eaten 
by  Dall's  porpoises  from  Monterey  Bay.  Pacific 
hake  have  been  reported  as  important  food  for 
Dall's  porpoise  (Scheffer  1953;  Norris  and  Pres- 
cott  1961;  Fiscus  and  Niggol  1965;  Loeb  1972). 
The  hakelike  fish  (Laemonema,  family  Moridae) 
occurs  in  the  diet  of  Dall's  porpoise  from  Jap- 
anese waters  (Wilke  and  Nicholson  1958). 

No  cephalopods  were  found  in  any  of  the  five 
Dall's  porpoise  stomachs  from  my  northern  Cal- 
ifornia sample. 

Delphinus  delphis,  Pacific  common  dolphin 

Common  dolphins  seldom  are  sighted  north  of 
the  California-Oregon  border  and  are  rare  be- 
yond the  100-fathom  (183-m)  line  (National 
Oceanic  and  Atmospheric  Administration  1974). 
Four  Delphinus  stomachs  collected  off  Califor- 
nia contained  fish  and  cephalopods  (Fiscus  and 
Niggol  1965).  One  female  had  unidentified  fish 
otoliths  and  another  stomach  contained  (by  vol- 
ume) 60  percent  squid,  25  percent  saury,  and  15 
percent  northern  anchovy.  One  male  dolphin 
taken  at  sea  contained  90  percent  Loligo  and  10 
percent  saury.  Another  stomach  contained  60 
percent  lanternfish  (Myctophidae)  and  40  per- 
cent squid  (Gonatus  sp.,  20  percent;  Onycho- 
teuthis sp.,  10  percent;  unidentified  squid,  10 
percent)  (Fiscus  and  Niggol  1965). 

Observations  in  California  waters  indicate 
that  common  dolphins  are  present  in  inshore 


waters  throughout  the  year  (Norris  and  Prescott 
1961).  The  major  foods  seem  to  be  sardines,  an- 
chovies, sauries,  small  bonito,  and  squid  (Norris 
and  Prescott  1961). 

Schmidt  (1923)  removed  15,191  otoliths  from 
the  stomach  of  one  Delphinus.  These  otoliths 
represented  five  species  of  small  fish  (7596  in- 
dividuals). Frost  (1924)  looked  at  4338  of  these 
same  otoliths  and  identified  six  species  in  three 
families.  Myctophid  fish  accounted  for  4324  of 
these  4338  otoliths.  The  fishes  represented  prob- 
ably did  not  constitute  a  "full"  meal  for  this 
dolphin  (Fitch  and  Brownell  1968). 

Many  common  dolphins  stranded  in  southern 
California  had  empty  stomachs  (Robert  Brow- 
nell, Jr.,  Smithsonian  Institution,  pers.  comm. 
[1970]).  Fitch  and  Brownell  (1968)  examined  two 
which  had  133  and  119  otoliths.  Anchovy  re- 
mains (141  sagittae)  were  the  most  abundant. 
One  dolphin  had  eaten  63  fishes  representing  six 
families.  Anchovy,  myctophids,  and  saury  were 
represented  in  the  other  common  dolphin  ex- 
amined. These  authors  speculated  that  both 
Lagenorhynchus  and  Delphinus  feed  on  meso- 
pelagic  fish  at  depths  exceeding  120  m  (Fitch  and 
Brownell  1968). 

The  Pacific  common  dolphin  (CAS  2340)  I  ex- 
amined had  11  otoliths  assigned  to  two  species; 
medusafish  (Icichthys  lockingtoni)  and  an  os- 
merid.  This  specimen  also  had  four  Loligo 
beaks.  Medusafish  are  most  abundant  around 
jellyfish  and  in  the  upper  150  ft  (ca.  46  m)  of  the 
ocean  (John  Fitch,  California  Dept.  of  Fish  and 
Game,  pers.  comm.  [1973]).  Apparently  this  dol- 
phin had  fed  near  the  surface. 


Lagenorhynchus  obliquidens, 

Pacific  white-sided  dolphin 

This  dolphin  has  received  careful  attention 
from  west  coast  biologists  during  the  last  25 
years  (Scheffer  1950b,  1953;  Brown  and  Norris 
1956;  Houck  1961;  Norris  and  Prescott  1961). 
These  authors  reported  sardine,  Pacific  herring, 
salmon,  northern  anchovy,  "scad"  (=jack- 
mackerel),  Pacific  saury,  squid,  and  jellyfish 
remnants  as  food  items  (Table  8). 

Large  feeding  aggregations  of  California  sea 
lions,  elephant  seals,  common  dolphins,  and  Pa- 
cific white-sided  dolphins  have  been  observed 
by  various  authors  (Norris  and  Prescott  1961; 
Fiscus  and  Niggol  1965).  Mixed  schools  of  com- 
mon and  white-sided  dolphins  have  been  noted 


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TABLE  8.     STOMACH  CONTENTS  OF  Lagenorhynchus  obliquiitens  FROM  THE  WEST  COAST  OF  NORTH  AMERICA  (CALIFORNIA). 


Locality 

Sex 

Date 

Source 

Type  of  contents 

w  Trinidad 

F 

11  Sep  1958 

(3) 

Pacific  sauries,  "scad" 

Humboldt  Co. 

M 

26  Dec.  1968 

(6) 

Pacific  sanddab.  Pacific  hake. 

eulachon,  squid 

Marin  Co. 

F 

9  Feb.  1970 

(6) 

no  ID  (fish) 

San  Mateo  Co. 

M 

29  Sep.  1972 

(6) 

shortbelly  rockfish,  northern 

anchovy,  plainfin  midshipman 

San  Mateo  Co. 

F 

6  May  1973 

(6) 

Pacific  hake,  jackmackerel 

w  Santa  Cruz 

M 

4  Mar.  1959 

(4) 

trace  squid 

w  Santa  Cruz 

F 

5  Mar.  1959 

(4) 

northern  anchovy.  Pacific  hake. 

squid 

Monterey  Co. 

F 

21  Apr.  1973 

(6) 

night  smelt 

Monterey  Co. 

F 

29  May  1970 

(6) 

squid,  octopus 

w  Pt.  Piedras 

F 

27  Feb.  1959 

(4) 

northern  anchovy,  squid 

NW  Morro  Bay 

M 

22  Feb.  1959 

(4) 

northern  anchovy.  Pacific  hake. 

squid 

w  Morro  Bay 

F 

14  Feb.  1959 

(4) 

trace  squid 

s  Anacapa  I. 

M 

27  Feb.  1952 

(1) 

jellyfish,  squid 

Santa  Monica 

9 

22  Aug.  1963 

(5) 

Pacific  hake,  northern  anchovy, 

white  seaperch,  cephalopods 

s  San  Pedro 

F 

6  June  1953 

(2) 

anchovies,  squid 

Long  beach 

F 

21  Aug.  1967 

(5) 

northern  anchovy,  Pacific  hake. 

cephalopods,  queenfish 

(1)  Scheffer  1953. 

(2)  Brown  and  Norris  1956. 

(3)  Houck  1961. 

(4)  Fiscus  and  Niggol  1965. 

(5)  Fitch  and  Brownell  1968. 

(6)  Current  study  1973. 


off  southern  California  where  the  white-sided 
dolphins  are  common,  but  north  of  San  Francis- 
co only  small  groups  of  Lagenorhynchus  have 
been  seen  (Fiscus  and  Niggol  1965). 

In  southern  California  waters,  white-sided 
dolphins  have  a  distinct  seasonal  movement  dur- 
ing the  summer  and  fall  which  correlates  with  a 
shift  from  anchovies  and  squids  as  a  principal 
food  to  the  offshore  schools  of  Pacific  saury. 
This  conclusion  is  from  field  observations  only 
and  from  one  stomach  analysis  (Brown  and  Nor- 
ris 1956).  No  seasonal  migration  of  white-sided 
dolphins  has  been  observed  in  northern  Califor- 
nia. A  migration  of  white-sided  dolphins  to 
northern  California  may  correlate  with  oceanic 
current  shifts  and/or  fish  migratory  patterns. 

Pacific  sanddab,  eulachon,  night  smelt,  short- 
belly  rockfish,  and  plainfin  midshipman  can  now 
be  added  to  the  known  species  of  food  fishes 
reported  from  white-sided  dolphins.  I  recovered 
89  otoliths-,  only  3  of  which  represent  open- 
water  fishes.  The  great  number  of  otoliths  from 
inshore  schooling  fishes  (Table  6)  tends  to  indi- 
cate that  this  dolphin  eats  abundant,  small  fishes 
(osmerids,  midshipman,  and  juvenile  rockfish). 


The  fact  that  white-sided  dolphins  feed  on  five 
kinds  of  cephalopods  gives  the  impression  that 
these  invertebrates  are  a  major  food  resource 
(Tables  5  and  9).  My  data  indicate  that  the  white- 
sided  dolphin  apparently  is  not  dependent  on 
cephalopods  in  northern  California;  only  three 
of  seven  stomachs  had  molluscan  remains.  One 
dolphin  (REJ  625)  contained  147  of  the  155  ceph- 
alopod  beaks  found. 

Grampus  griseus,  Risso's  dolphin 

Until  recent  stranding  records  were  published 
(Orr  1966;  Paul  1968;  Stroud  1968;  Hatler  1971) 
this  dolphin  was  known  only  from  the  type  of 
Grampus  stearnsii  (Dall)  collected  at  Monterey 
Bay  in  1873.  All  four  previous  specimens  re- 
ported were  males,  and  only  two  contained  iden- 
tifiable food  remains.  Orr  (1966)  reported  a  Ris- 
so's dolphin  from  San  Mateo  County  that 
contained  two  pairs  of  beaks  from  the  squid 
Dosidicus  gigas.  Stroud  (1968)  listed  seven  cat- 
egories of  cephalopod  beaks  from  a  male  Risso's 
dolphin  recovered  in  Washington:  Onychoteu- 
this  (1),  Octopodoteuthis  (1),  Chiroteuthis  (16), 
Gonatus  (4),  Gonatidae — form  A  (7),  Gonati- 


JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


427 


dae — form  B  (13),  and  unidentified  beaks  (2).  No 
fishes  have  been  found  in  any  Risso's  dolphin. 
I  examined  a  single  specimen  at  Southeast 
Farallon  Island  on  20  May  1973.  All  standard 
measurements  (275  cm  total  length)  were  taken, 
and  the  reproductive  tract,  blood  sample,  and 
complete  skeleton  were  saved.  No  parasites 
were  discovered,  but  a  goose-neck  barnacle 
(Pollicipes  polymerus)  and  a  single  hydroid 
(Aglaophenia  latirostris)  were  in  the  stomach. 
Orr  (1951)  theorized  that  such  material  is  swal- 
lowed incidentally  as  the  animal  thrashes  close 
to  shore.  The  barnacle  and  hydroid  reported 
here  were  probably  accidentally  ingested.  Both 
kinds  of  invertebrates  are  plentiful  on  the  inter- 
tidal  shores  of  the  Farallon  Islands.  Hatler 
(1971)  reported  that  plant  material  seems  to  be 
an  "herbal  remedy"  in  Grampus.  Risso's  dol- 
phin is  probably  an  invertebrate  feeder  like 
Globicephala,  which  eats  only  squid  as  reported 
by  Sergeant  and  Fisher  (1957).  Grampus  lack 
teeth  in  the  upper  jaw  and  may  feed  solely  on 
soft  invertebrates. 

Kogia  simus,  Dwarf  sperm  whale 

Brownell  obtained  three  dwarf  sperm  whales 
from  Japanese  waters.  Although  a  good  com- 
parative collection  of  Japanese  fishes  was  lack- 
ing, Fitch  and  Brownell  (1968)  presented  a  table 
indicating  that  18  different  species  of  fish  in  7 
families  were  in  the  stomachs.  Because  two 
families  (Macrouridae  and  Moridae)  are  inhab- 
itants of  deep  water,  these  authors  speculated 
that  Kogia  feeds  800  ft  (244  m)  or  more  beneath 
the  surface.  Other  authors  have  stated  that  most 
specimens  of  Kogia  had  eaten  cephalopods  or 
pelagic  crustaceans  (National  Oceanic  and  At- 
mospheric Administration  1974). 

Scheffer  and  Slipp  (1948)  examined  a  male 
pygmy  sperm  whale  specimen  from  Washington. 
It  had  500  cc  of  nematodes  and  food  fragments 
in  its  stomach;  11  eye  lenses  (including  5  from 
squid),  15  squid  beaks,  21  otoliths  of  an  uniden- 
tified fish,  1  crab  limb,  maxillary  bones  of  2 
specimens  of  Trichodon  (?),  and  fragments  of 
shrimp  Pasiphaea,  Pandalus,  Pandalopsis. 

Kogia  has  a  discontinuous  distribution  and  is 
rather  poorly  known  from  the  west  coast.  Few 
have  been  seen  alive.  Beach-cast  specimens 
have  provided  material  for  anatomical,  taxo- 
nomic,  and  distributional  studies,  but  essentially 
nothing  is  known  of  its  life  history  (Handley 
1966). 


Because  of  difficulty  of  identification  prior  to 
Handley's  work  (1966),  most  earlier  literature 
could  apply  to  either  the  pygmy  sperm  whale 
(K.  breviceps)  or  the  dwarf  sperm  whale.  Only 
in  recent  studies  can  identification  be  trusted. 

One  K.  simus  recovered  at  Thorton  Beach, 
San  Mateo  County,  by  the  California  Academy  of 
Sciences  (CAS  2382,  male,  204  cm  total  length) 
contained  217  beaks  representing  1 12  individuals 
of  these  families:  Octopoteuthidae,  Onycho- 
teuthidae,  Enoploteuthidae,  Histioteuthidae, 
Gonatidae,  Chiroteuthidae.  In  addition,  a  single 
pair  of  otoliths  in  its  stomach  was  from  a  plainfin 
midshipman  (Porichthys).  This  is  the  most 
northern  record  of  Kogia  simus  along  the  west 
coast. 

DISCUSSION 

Statements  about  marine-mammal  diets  fre- 
quently have  been  vague  and  misleading  and 
give  a  false  impression  of  the  role  these  animals 
play  in  the  marine  ecosystem.  Much  more  in- 
formation is  needed  to  understand  predator-prey 
relationships  in  California  waters  (Steele  1970). 
California  waters  are  rich  in  commercially  im- 
portant fishes,  and  these  fishes  are  well  studied. 
However,  it  is  also  pertinent  to  consider  the 
stocks  of  noncommercial  fishes  and  the  dynam- 
ics of  such  populations.  The  distribution  of  both 
predator  and  prey,  including  the  availability  of 
the  latter  throughout  the  year,  is  of  prime  im- 
portance. 

Adult  marine  mammals  are  usually  migratory. 
The  causes  of  these  movements  are  largely  un- 
known, but  to  some  extent  marine  mammals  re- 
spond to  the  seasonal  abundance  of  food.  The 
breeding  cycle  and  its  influence  on  food  gath- 
ering have  not  been  studied.  The  smaller  whales 
are  not  associated  with  breeding  rookeries  as  are 
pinnipeds.  Cetaceans  constantly  search  for  prey 
and  apparently  lack  the  feeding-nursing  cycle 
characteristically  found  in  seals  and  sea  lions. 

The  behavior  and  feeding  techniques  of  all 
marine  mammals  are  poorly  documented.  Es- 
cape strategies  of  prey  species  are  not  well 
known.  The  sheer  abundance  of  smaller  prey 
items  allows  the  escape  of  some  individuals  from 
predators.  Some  fishes  and  invertebrates  reduce 
predation  upon  themselves  by  camouflage  or 
disguise.  Armor  and  spines  must  also  help  some 
prey  to  reduce  losses.  Representatives  of  the 
following  common  families  of  fishes  were  com- 
pletely absent  from  the  marine  mammal  stom- 


428 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  16 


TABLE  9.     STOMACH  CONTENTS  OF  MARINE  MAMMALS  FOUND  IN  THIS  STUDY  (volume  of  dry  fish  hones,  cephalopod 
identification,  and  miscellaneous  items). 


Field  number 

Fish  bones 
(cc) 

Cephalopod  beaks 

Upper 

Lower 

Identification 

Zalophus 

68-40 

950 

_ 

- 

— 

317  LGB 

400 

- 

_ 

— 

647 

35 

2 

3 

Octopus  sp. 

660 

4000 

- 

1 

Octopus  sp. 

662 

trace 

3 

- 

L.  opalescens 

664 

250 

- 

- 

— 

665 

30 

23 

17 

L.  opalescens 

666 

5 

- 

- 

— 

667 

trace 

- 

- 

— 

685 

120 

- 

- 

— 

1  —  Polinicex  shell, 

Ulva,  eelgrass 

686 

820 

- 

- 

— 

692 

45 

- 

1 

Octopus  sp. 

244 

- 

1 

3 

L.  opalescens 

Eumetopias 

69-8 

600 

_ 

1 

Chiroteuthis 

247 

- 

1 

1 

L.  opalescens 

249 

- 

6 

7 

L.  opalescens 

629 

- 

2 

2 

Octopus  sp. 

637 

2850 

2 

2 

L.  opalescens 

655 

1  —  rock 

- 

1 

Onychoteuthis 

675 

trace 

_ 

_ 

— 

677 

5  —  rocks,  trace 

- 

- 

— 

688 

240 

- 

- 

— 

699 

3200 

- 

1 

Octopus  sp. 

Phocn 

69-24 

trace 

_ 

_ 

— 

646 

40 

2 

1 

Octopus  sp. 

671 

10 

- 

- 

— 

696 

400 

10 

3 

Octopus  sp. 

Callorhinu.s 

212 

7  —  external  fish 

- 

- 

— 

isopods  (Riftgia  ?) 

243 

- 

7 

7 

L.  opalescens 

Delphinus 

CAS  2340 

3 

1 

1 

L.  opalescens 

Lagenorhynchus 

HSC  68-9 

- 

1 

- 

Ahrtiliopsis 

237 

- 

1 

1 

Octopoteuthis 

1 

3 

L.  opalescens 

- 

1 

unknown  Gonatidae 

625 

- 

1 

1 

Onychoteuthis 

79 

66 

L.  opalescens 

- 

1 

Gonatus  sp. 

652 

trace 

- 

- 

— 

Phocoena 

241 

- 

9 

7 

L.  opalescens 

450 

15 

35 

17 

L.  opalescens 

653 

- 

8 

5 

L.  opalescens 

661 

- 

- 

1 

L.  opalescens 

673 

75 

1 

1 

L.  opalescens 

JONES:  FOOD  HABITS  OF  SMALLER  MARINE  MAMMALS 


429 


CAS  2398 


TABLE  9.     CONTINUED. 


Field  number 

Cephalopod  beaks 

Identification 

cc                                    Upper                 Lower 

HSC  73-4 
CAS  2384 
CAS  2385  (476) 
CAS  2392 

180 
2                          3 
trace 
35                        36 

L.  opalescens 
L.  opalescens 

plus  13  whole  Loligo 
Morotheuthis  sp. 


achs  examined  in  this  study:  Cottidae,  Agoni- 
dae,  Serranidae,  Blenniidae,  Clinidae,  and 
Scombridae.  These  marine  fishes  are  probably 
detected  and  perhaps  discriminated  as  nonprey 
items.  Possibly  these  fishes  possess  mechanisms 
to  escape. 

Sick  or  injured  marine  mammals  will  starve 
rapidly.  Animals  not  showing  obvious  causes  of 
death  presumably  were  sick  or  injured  and  sel- 
dom had  anything  in  their  stomachs.  Specimens 
showing  evidence  of  violent  sudden  death  had 
intact  squid  or  fish.  Others  had  only  digestion- 
resistant  items  such  as  beaks  or  otoliths.  One 
harbor  porpoise  (CAS  2390)  had  swallowed  a  46- 
cm  Pacific  hake,  the  anterior  end  of  which  was 
partially  digested.  No  food  item  in  this  study 
showed  evidence  of  having  been  chewed  or  cut 
by  the  consumer.  Field  observations  indicate 
that  food  is  torn  apart  by  much  head  shaking, 
and  the  teeth  only  aid  in  the  capture  and  holding 
of  prey.  The  lack  of  specialized  forelimbs  with 
which  to  manipulate  food  is  evident  in  the  swal- 
lowing of  whole  food. 

Sexual  dimorphism  in  the  size  of  pinnipeds 
(Scheffer  1958)  should  be  reflected  in  feeding 
rates  and  the  species  or  size  of  prey  selected, 
but  no  published  data  are  available  to  substan- 
tiate this.  However,  such  resource  partitioning 
is  shown  between  species  (Table  6). 

It  appears  that  prey  selection  in  marine  mam- 
mals is  specialized,  and  this  conclusion  is  sup- 
ported by  the  available  data.  Comparison  of  the 
food  of  Phoca  with  that  of  Eumetopias  and  Zal- 
ophus  (Tables  1 ,  4,  and  7)  shows  less  dependence 
on  pelagic  fishes  by  Phoca.  Perch,  eelpout,  and 
greenling  (80  percent  of  the  fish  eaten  by  Phoca) 
are  typically  shallow-bottom  species  which  live 
near  rocky  habitats.  Zalophus  characteristically 
feeds  on  Pacific  hake,  northern  anchovy,  and 
rockfish  (93  percent  of  the  fish).  These  open- 


water  fishes  are  very  abundant  (Ahlstrom  1965; 
Bell  (1971).  Eumetopias,  while  also  using  Pa- 
cific hake  and  rockfish,  relied  more  heavily  on 
bottom-dwelling  flatfish  and  cusk-eel  (29. 1  per- 
cent of  the  fish).  Octopus  was  the  only  cepha- 
lopod  found  in  the  stomachs  of  harbor  seals,  al- 
though several  other  cephalopods  are  taken  by 
California  and  northern  sea  lions  (Table  9). 
Stomachs  of  Eumetopias  frequently  had  stones 
in  them  (Table  9).  Apparently  there  is  some  se- 
lection for  these  nonfood  items  by  these  sea 
lions.  The  depth  of  the  sea  at  which  these  pin- 
nipeds feed  is  unknown.  All  prey  items  normally 
eaten  by  Eumetopias  inhabit  water  less  than 
about  200  m  deep.  Thus  Zalophus  feeds  on 
schooling  fishes  while  Eumetopias  feeds  on  bot- 
tom fishes  (Table  6). 

The  mean  feeding  rate  of  small  cetaceans  as 
described  by  Sergeant  (1969)  is  10.8  percent  of 
their  body  weight  per  day.  No  comparative  in- 
formation is  available  on  feeding  rates  of  juve- 
nile individuals,  nor  have  sexual  or  seasonal 
differences  in  feeding  rates  been  published. 
Captive  Arctic  seals  (Cystophora,  Pagophilus) 
require  food  in  amounts  of  3-5  percent  of  their 
body  weight  per  day  in  order  to  maintain  good 
health  (Blix  et  al.  1973).  Daily  food  consump- 
tion of  fur  seals  (10  percent  of  body  weight), 
northern  sea  lion  (4  percent  of  body  weight), 
and  harbor  seal  (1 1  percent  of  body  weight)  are 
recorded  on  field-collected  specimens  by  Spald- 
ing  (1964).  These  data  do  not  include  informa- 
tion on  body  size,  reproductive  state,  or  activity 
factors  which  influence  the  food  consumption 
of  these  pinnipeds. 

The  smaller  odontocetes  fed  on  more  cepha- 
lopods (17  percent  of  the  diet)  than  did  the  pin- 
nipeds (5  percent)  (Tables  5  and  9).  The  diet  of 
the  sea  lions  in  this  study  was  95  percent  fish, 
compared  with  83  percent  in  the  cetaceans. 


430 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42.  No.  16 


Phocoena  fed  heavily  on  rockfish,  anchovy,  and 
juvenile  Pacific  hake.  Most  of  these  fishes  are 
semi-pelagic,  small,  and  probably  occur  near  the 
surface.  The  recoveries  of  harbor  porpoises 
were  limited  to  the  period  April  to  September 
and  therefore  do  not  reflect  a  year-round  dietary 
sample.  These  porpoises  prey  extensively  on 
Loligo,  which  is  present  throughout  the  year. 

Lagenorhynchus  associates  with  many  other 
species  of  marine  animals  and  perhaps  feeds  on 
a  wide  variety  of  prey.  Data  from  this  study 
(Table  8)  indicate  that  white-sided  dolphins  are 
generalized  feeders.  Osmerids  and  midshipmen, 
representing  two  distinct  habitats  (inshore 
schooling  and  bottom-dwelling),  were  major 
food  items.  Although  midshipmen  live  from  the 
intertidal  regions  to  a  depth  of  170  fathoms 
(about  311  m),  they  also  leave  the  bottom  in 
search  of  food  (Fitch  and  Lavenberg  1971).  We 
do  not  know  where  or  when  this  fish  is  eaten, 
but  it  is  an  important  component  in  the  diets  of 
all  marine  mammals  studied. 

ACKNOWLEDGMENTS 

So  many  individuals  have  contributed  to  this 
study  that  it  is  impractical  to  list  them  all.  The 
telephone  calls  from  federal,  state,  and  county 
officials  and  other  interested  persons  reporting 
carcasses  are  deeply  appreciated.  Personnel  at 
the  Point  Reyes  Bird  Observatory  deserve  spe- 
cial thanks.  The  facilities  of  these  institutions 
were  used:  California  Academy  of  Sciences; 
Museum  of  Vertebrate  Zoology,  University  of 
California,  Berkeley;  Humboldt  State  Univer- 
sity. 

Personnel  of  each  institution  extended  aid, 
especially  my  dissertation  committee  at  Hum- 
boldt State  University,  chaired  by  Dr.  W.  J. 
Houck.  The  patience  showed  by  the  manuscript 
reviewers  is  appreciated;  thank  you,  Drs.  S. 
Harris,  J.  Waters,  and  W.  Z.  Lidicker,  Jr.  The 
assistance  of  Mr.  John  E.  Fitch,  Research  Di- 
rector, State  Fisheries  Laboratory,  California 
Department  of  Fish  and  Game,  was  essential  for 
the  identification  offish  otoliths.  Mr.  Clifford  H. 
Fiscus,  National  Marine  Fisheries  Service,  Se- 
attle, Washington,  aided  with  the  identification 
of  the  cephalopod  beaks. 

My  able  field  assistants  located  as  many  old 
carcasses  as  I  did.  Thanks,  Pica,  Chepen, 
Dusty,  and  Ginger.  And  lastly,  the  support  of 
Lise  made  all  the  tasks  easier. 


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CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  17,  pp.  435^42,  21  figs. 


October  26,  1981 


STUDIES  ON  THE  NEBRIINI  (COLEOPTERA:  CARABIDAE), 

IV.  FOUR  NEW  NEBR1A  TAXA  FROM 

WESTERN  NORTH  AMERICA 

By 

David  H.  Kavanaugh 

Department  of  Entomology,  California  Academy  of  Sciences, 
Golden  Gate  Park,  San  Francisco,  California  94118 


ABSTRACT:  This  paper,  fourth  of  a  series  on  the  nebriine  carabid  beetles,  provides  names  for  four  undescribed 
taxa  from  western  North  America,  including:  Nebria  danmanni  (type-locality:  Deception  Basin,  Olympic  Na- 
tional Park,  Washington),  N.  sonorae  (Chipmunk  Flat,  Tuolumne  County,  California),  V.  turmaduodecima 
(Caribou  Basin,  Siskiyou  County,  California),  and  N.  meanyi  giulianii  (Milner  Creek,  Mono  County,  Califor- 
nia). Diagnostic  combination  of  characters  and  comment  on  geographical  distribution  are  provided  for  each 
new  taxon  and  distinguishing  features  are  illustrated. 


INTRODUCTION 

As  part  of  an  ongoing  project  on  the  Nebriini 
of  the  world,  Kavanaugh  (1979)  provided  names 
for  5  new  species  and  23  new  subspecies  of  ge- 
nus Nebria  Latreille  from  North  America  and 
updated  nomenclature  for  Nearctic  members  of 
the  genus.  Since  the  appearance  of  that  paper, 
three  new  species  and  one  new  subspecies  of 
Nebria  have  been  discovered. 

The  purpose  of  this  report,  an  addendum  to 
Kavanaugh  (1979),  is  to  provide  names  for  these 
new  taxa.  Names  are  needed  immediately  for 
use  in  several  other  manuscripts  and  by  other 
workers.  To  this  end,  data  and  discussion  pre- 
sented for  each  name  are  limited  to  little  more 
than  the  minimum  required  by  the  International 
Code  of  Zoological  Nomenclature  for  availabil- 
ity. Additional  information  on  all  Nearctic  Ne- 
bria taxa,  including  those  presented  here  as 
new,  will  be  presented  in  a  subsequent  paper 
now  in  preparation. 


MATERIALS 

This  study  is  based  on  examination  of  187 
adult  Nebria  specimens.  Following  is  a  list  of 
abbreviations  used  in  the  text  which  refer  to  the 
collections  from  which  specimens  were  received 
and/or  in  which  paratype  specimens  have  been 
deposited.  Names  of  curators  who  sent  speci- 
mens are  also  included. 


CAS — California  Academy  of  Sciences,  San  Francisco,  Cali- 
fornia 941 18;  D.  H.  Kavanaugh. 

CDA — California  State  Department  of  Food  and  Agriculture, 
Sacramento,  California  95814;  F.  G.  Andrews. 

DMan — D.  Mann.  University  of  Washington.  Seattle,  Wash- 
ington 98195. 

UASM — University  of  Alberta,  Strickland  Museum,  Edmon- 
ton, Alberta  T6G  2E3;  G.  E.  Ball. 

UCD— University  of  California,  Davis,  California  95616:  R. 
O.  Schuster. 

USNM — United  States  National  Museum  of  Natural  History, 
Smithsonian  Institution,  Washington,  D.C.  20560;  T.  L. 
Erwin. 


[435] 


436 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  17 


FIGURES  1-4.  Figs.  1-3.  Pronotum,  dorsal  aspect;  scale  line  =  1.0  mm.  1.  Nebria  lurmaduodecima  n.sp.  (Upper  Caribou 
Lake,  California).  2.  Nebria  clanintinni  n.sp.  (Deception  Basin,  Washington).  3.  Nebria  sonorae  n.sp.  (Chipmunk  Flat,  Cali- 
fornia). Fig.  4.  Head  and  pronotum,  dorsal  aspect,  Nebria  meanyi  giulianii  n.ssp.  (Milner  Creek,  California). 


METHODS 

Methods  which  relate  specifically  to  data  and 
results  presented  in  this  paper  (including  dissec- 
tion techniques  and  criteria  for  ranking  taxa)  are 
described  in  Kavanaugh  (1979).  A  broader  and 
more  detailed  discussion  of  preparative  and  pro- 
cedural methods  used  will  be  included  in  a  sub- 
sequent paper  (Kavanaugh,  manuscript  in  prep- 
aration). 

NEW  NEBRIA  TAX  A 

The  order  of  presentation  of  new  taxa  follows 
a  new  classification  of  Nearctic  Nebria  to  be 
presented  at  a  later  date  (Kavanaugh,  manu- 
script in  preparation). 

Nebria  turmaduodecima,  new  species 

(Figures  1,  5,  9,  13,  17,  21) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  California, 
Siskiyou  Co.,  Trinity  Alps,  Caribou  Basin  (south  rim),  at 


snowfield  edges,  2290m,  12  Aug.  1980,  Stop  #80-27  D.  H. 
Kavanaugh  collector"/" D.  H.  Kavanaugh  Collection"  [or- 
ange label]/"Holotype  Nebria  turmaduodecima  Kavanaugh 
del.  D.  H.  Kavanaugh  1981"  [red  label]/"California  Academy 
of  Sciences  Type  No.  13729."  PARATYPES:  140(65  males,  75 
females),  deposited  in  CAS,  CDA,  UASM,  UCD,  USNM. 

TYPE-LOCALITY. — Caribou  Basin,  2290  m,  Trinity  Alps, 
Siskiyou  County,  California. 

DIAGNOSTIC  COMBINATION. — Size  medium, 
standardized  body  length  of  male  less  than  12.0 
mm,  of  female  less  than  12.4  mm;  head  dark, 
with  pair  of  pale  spots  present  on  vertex  (par- 
tially fused  medially  in  some  individuals);  elytra 
slightly  shiny  (microsculpture  moderately  im- 
pressed, meshes  isodiametric  or  very  slightly 
transverse),  without  metallic  reflection  (very 
slightly  developed  in  a  few  individuals  only); 
pronotum  (Fig.  1)  with  midlateral  and  basolater- 
al  setae  present;  elytral  silhouette  markedly 
ovoid,  narrowed  basally  (Fig.  5);  hindwing  (Fig. 
9)  vestigial;  median  lobe  of  male  genitalia  as  in 


KAVANAUGH:  NEW  NEBRIA  TAXA 


437 


FIGURES  5-8.  Basal  region  of  left  elytron,  dorsal  aspect;  scale  line  =  1.0  mm.  5.  Nehriu  turmiuluodec'una  n.sp.  (Upper 
Caribou  Lake,  California).  6.  Nebria  danmunni  n.sp.  (Deception  Basin,  Washington).  7.  Nebria  sonorue  n.sp.  (Chipmunk  Flat, 
California).  8.  Nebria  ineanyi  giulianii  n.ssp.  (Milner  Creek,  California). 


Figure  13;  bursa  copulatrix  of  female  as  in  Fig- 
ure 17;  specimen  from  Trinity  Alps,  northwest- 
ern California. 

DERIVATION  OF  TAXON  NAME. — The  species 
epithet  is  a  combination  of  the  Latin  words  for 
"troop"  (=turma)  and  "twelve"  ^duodeci- 
mo). It  is  a  pleasure  for  me  to  name  this  species 
in  honor  of  the  boys  of  Troop  12,  Boy  Scouts  of 
America,  Petaluma,  California,  who  assisted  me 
in  collecting  the  first  known  specimens  of  this 
species. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  21. 
Known  only  from  Caribou  Basin  in  the  Trinity 
Alps  of  northwestern  California;  range  probably 
restricted  to  the  Trinity  Alps.  I  have  studied 
specimens  from  the  following  localities. 

United  States  of  America 

CALIFORNIA:  Siskiyou  County,  Caribou  Basin  (south  rim 
[2290  m])  [Aug.]  (45;  CAS),  Upper  Caribou  Lake  (east  shore 
[2100-2130  m])  [Aug.]  (96;  CAS,  CDA,  UASM,  UCD, 
USNM). 


Nebria  danmanni,  new  species 

(Figures  2,  6,  10,  14,  18,  21) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  Washing- 
ton, Olympic  National  Park,  Deception  Basin,  6000ft.,  4  Sept. 
1976  D.  Mann  collector'T'Holotype  Nebria  danmanni  Ka- 
vanaugh  det.  D.  H.  Kavanaugh  1981"  [red  label]/"California 
Academy  of  Sciences  Type  No.  13730."  PARATYPES:  28  (13 
males,  15  females),  deposited  in  CAS,  DMan,  UASM, 
USNM. 

TYPE-LOCALITY. — Deception  Basin,  1830  m,  Olympic  Na- 
tional Park,  Washington. 

DIAGNOSTIC  COMBINATION. — Dorsal  surface 
very  shiny,  elytral  microsculpture  formed  of 
markedly  transverse  meshes;  pronotum  (Fig.  2) 
with  apical  angle  markedly  projected  anteriorly, 
basal  angle  rectangular,  denticulate,  and  mod- 
erately projected  posteriorly,  basal  sinuation  of 
lateral  margin  very  long  and  shallow,  midlateral 
seta  absent;  elytral  silhouette  subrectangular, 
markedly  elongate,  slightly  narrowed  basally, 
humeral  angle  (Fig.  6)  markedly  distinct,  hu- 
meral carina  markedly  developed  and  projected 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  17 


FIGURES  9-12.  Left  hindwing;  scale  line  =  1.0  mm.  9.  Nehria  turmaduodecima  n.sp.  (Upper  Caribou  Lake,  California). 
10.  Nebrin  tlannninni  n.sp.  (Deception  Basin,  Washington).  1 1.  Nehria  sonorae  n.sp.  (Chipmunk  Flat,  California).  12.  Nebria 
ineanyi  giulianii  n.ssp.  (Milner  Creek,  California). 


anteriorly;  hindwing  (Fig.  10)  markedly  short- 
ened and  narrowed;  middle  tibia  moderately 
concave  to  slightly  sulcate  dorsally  at  middle, 
brush  of  dorsal  setae  moderately  dense  subapi- 
cally;  hindcoxa  bi-  or  plurisetose  basally;  third 
to  fifth  visible  abdominal  sterna  each  with  two 
or  more  pairs  of  posterior  paramedial  setae;  me- 
dian lobe  of  male  genitalia  as  in  Figure  14;  bursa 
copulatrix  of  female  as  in  Figure  18. 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  species  in  honor  of  my 
friend  and  field  companion,  Daniel  H.  Mann, 
who  collected  the  first  known  specimens,  in- 
cluding the  holotype,  of  this  species. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  21. 
Known  only  from  Deception  Basin  in  south- 
eastern Olympic  National  Park,  Washington; 
range  probably  restricted  to  the  Olympic  Moun- 
tains at  high  elevations.  I  have  studied  speci- 
mens from  the  following  localities. 

United  States  of  America 

WASHINGTON:  Olympic  National  Park,  Deception  Basin 
([1830  m])  [Sep.]  (II;  CAS,  DMan).  Mount  Mystery  (east 
slope  [1800-1860  m])  [July]  (18;  CAS,  USNM). 


Nebria  sonorae,  new  species 

(Figures  3,  7,  11,  15,  19,  21) 

HOLOTYPE,  a  male,  in  CAS  (on  indefinite  loan  deposit  from 
UCD)  labelled:  "Chipmunk  Flat  Tuolumne  Co.,  Calif.  VIII-9- 
60"/"R.  R.  Montanucci  Collector'VHolotype  Nebria  sono- 
rae Kavanaugh  det.  D.  H.  Kavanaugh  1981"  [red  label]/"Cal- 
ifornia  Academy  of  Sciences  Type  No.  13731."  PARATYPES: 
two  females,  deposited  in  CAS,  UCD. 

TYPE-LOCALITY. — Chipmunk  Flat,  Tuolumne  County,  Cal- 
ifornia. 

DIAGNOSTIC  COMBINATION. — Dorsal  surface 
moderately  shiny,  elytral  microsculpture  mod- 
erately impressed,  formed  of  isodiametric  (or 
nearly  so)  meshes;  pronotum  (Fig.  3)  with  apical 
angle  moderately  projected,  basal  angle  rectan- 
gular or  slightly  acute,  not  projected  posteriorly, 
basal  sinuation  of  lateral  margin  moderate  in 
length  and  depth,  midlateral  seta  absent;  elytral 
silhouette  subrectangular,  moderately  elongate, 
moderately  narrowed  basally,  humeral  angle 
(Fig.  7)  moderately  distinct,  humeral  carina  well 
developed  and  projected  anteriorly;  hindwing 
(Fig.  1 1)  full-sized  and  width;  middle  tibia  mod- 
erately sulcate  dorsally  at  middle,  brush  of  dor- 
sal setae  moderately  dense  subapically;  hind 


KAVANAUGH:  NEW  NEBRIA  TAXA 


439 


FIGURES  13-16.  Median  lobe  of  male  genitalia,  left  lateral  aspect;  scale  line  =  1.0  mm.  13.  Nebria  turmaduodecima  n.sp. 
(Upper  Caribou  Lake,  California).  14.  Nehriu  ilanmanni  n.sp.  (Deception  Basin,  Washington).  15.  Nebria  sonorae  n.sp. 
(Chipmunk  Flat,  California).  16.  Nebriu  meanyi  giulianii  n.ssp.  (Montgomery  Creek,  California). 


coxa  bi-  or  plurisetose  basally;  third  to  fifth  vis- 
ible abdominal  sterna  each  with  two  or  more 
pairs  of  posterior  paramedial  setae;  median  lobe 
of  male  genitalia  as  in  Figure  15;  bursa  copula- 
trix  of  female  as  in  Figure  19. 

DERIVATION  OF  TAXON  NAME. — This  species 


is  named  for  Sonora  Pass,  which  is  the  low  point 
on  the  main  divide  of  the  Sierra  Nevada  between 
Tuolumne  and  Mono  counties  and  is  the  domi- 
nant landmark  in  the  vicinity  of  the  type-locali- 
ty. 
GEOGRAPHICAL  DISTRIBUTION. — Figure  21. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  17 


FIGURES  17-18.  Bursa  copulatrix  (a,  dorsal  aspect;  b,  left  lateral  aspect;  c,  mid-sagittal  outline,  left  lateral  aspect):  scale 
line  =  1.0  mm.  17.  Nebria  turmaduodecima  n.sp.  (Upper  Caribou  Lake,  California).  18.  Nehria  dcinmcinni  n.sp.  (Deception 
Basin,  Washington). 


Known  only  from  Chipmunk  Flat,  Tuolumne 
County,  California,  and  the  stream  (southeast  of 
and  above  the  Flat)  which  drains  the  northwest 

flank  Of  Leavitt  Peak  massif;  range  probably  re- 

stricted  to  that  portion  of  the  Sierra  Nevada  be- 


tween  Sonora  Pass  and  Buckeye  Pass.  I  have 
studied  specimens  from  the  following  localities. 
united  states  of  America 

CALIFORNIA:   Tuolumne  County,  Chipmunk   Flat   (and 

stream  SE  [2440-2680  m])  [June,  Aug.]  (3;  CAS,  UCD). 


KAVANAUGH:  NEW  NEBRIA  TAXA 


441 


FIGURES  19-20.  Bursa  copulatrix  (a,  dorsal  aspect;  b,  left  lateral  aspect;  c,  mid-sagittal  outline,  left  lateral  aspect);  scale 
line  =  1.0  mm.  19.  Nebria  sonorae  n.sp.  (Chipmunk  Flat,  California).  20.  Nebria  meanyi  giulianii  n.ssp.  (Milner  Creek, 
California). 


Nebria  meanyi  giulianii,  new  subspecies 

(Figures  4,  8,  12,  16,  20,  21) 

HOLOTYPE,  a  male,  in  CAS,  labelled:  "U.S.A.,  California, 
Mono  County,  White  Mts.,  Montgomery  Creek,  2380m,  21 
June  1980  D.  Giuliani  collector'T'Holotype  Nebria  meanyi 
giulianii  Kavanaugh  del.  D.  H.  Kavanaugh  1981"  [red  label]/ 
"California  Academy  of  Sciences  Type  No.  13732."  PARA- 
TYPES:  13  (3  males,  10  females),  deposited  in  CAS. 

TYPE-LOCALITY. — Montgomery  Creek,  2380  m,  Mono 
County,  California. 

DIAGNOSTIC  COMBINATION. — Size  medium, 
standardized  body  length  of  male  less  than  12.0 


mm,  of  female  less  than  12.3  mm;  head  dark, 
with  pair  of  pale  spots  present  on  vertex;  elytra 
with  slight  to  moderate  metallic  blue,  green,  or 
violet  reflection;  head  (Fig.  4)  relatively  broad, 
antennal  scape  short,  cylindrical;  pronotum 
(Fig.  4)  relatively  short,  wide,  moderately 
cordate,  slightly  broad  basally,  hind  angle  rec- 
tangular, moderately  projected  posteriorly,  mid- 
lateral  seta  present;  elytral  silhouette  subrec- 
tangular,  broad  basally,  basal  margination  (Fig. 
8)  moderately  concave;  hindwing  (Fig.  12)  full- 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  17 


sized;  median  lobe  of  male  genitalia  as  in  Figure 
16;  bursa  copulatrix  of  female  as  in  Figure  20; 
specimen  from  White  Mountains  of  eastern  Cal- 
ifornia (Fig.  21). 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  subspecies  in  honor  of 
Derham  Giuliani,  a  naturalist  and  friend  whose 
long-term  interest  has  been  the  exploration  and 
biotic  inventory  of  remote  and  poorly  sampled 
parts  of  California  and  Nevada.  On  one  of  his 
recent  forays,  he  discovered  populations  of  this 
new  subspecies. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  21. 
Known  only  from  two  localities  on  the  western 
slope  of  the  White  Mountains  of  eastern  Cali- 
fornia; range  probably  restricted  to  that  moun- 
tain chain.  I  have  studied  specimens  from  the 
following  localities. 

United  States  of  America 

CALIFORNIA:  Mono  County,  Milner  Creek  ([2440  m])  [July] 
(9;  CAS),  Montgomery  Creek  ([2380  m])  [June]  (5;  CAS). 

ACKNOWLEDGMENTS 

In  addition  to  those  individuals  whose  contri- 
butions I  acknowledged  in  naming  three  of  the 
taxa  described  above,  I  thank  Terry  L.  Erwin 
and  my  son,  Thomas  W.  Kavanaugh,  for  their 
help  and  companionship  during  field  work  in  the 
Olympic  Mountains  (Washington)  and  Sonora 
Pass  area  (California),  respectively. 

LITERATURE  CITED 

KAVANAUGH,  D.  H.  1979.  Studies  on  the  Nebriini  (Coleop- 
tera:  Carabidae),  III.  New  Nearctic  Nebria  species  and  sub- 
species, nomenclature!  notes,  and  lectotype  designations. 
Proc.  Calif.  Acad.  Sci.  42:87-133. 


FIGURE  21.  Map  of  geographical  distributions:  Nehria  tur- 
mutluoilecinui  n.sp.  =  solid  circle;  Nehrici  diinmunni  n.sp.  = 
solid  triangle;  Nebria  soiwrae  n.sp.  =  solid  star;  Nehrici 
ineanyi  giulianii  n.ssp.  =  solid  square. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  941 18 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  18,  pp.  443-454,  4  figs. 


May  14,  1982 


DENDROGASTER  (CRUSTACEA:  ASCOTHORACIDA)  FROM 

CALIFORNIA:  SEA-STAR  PARASITES  COLLECTED 

BY  THE  ALBATROSS 

By 
Mark  J.  Grygier 

Scripps  Institution  of  Oceanography,  A-008,  University  of  California,  San  Diego, 
La  JoUa,  California  92093 


ABSTRACT:  Californian  specimens  of  the  ascothoracid  crustacean  genus  Dendrogaster,  discovered  by  W.  K. 
Fisher  in  sea-stars  collected  by  the  ALBATROSS,  are  reviewed.  Dendrogaster  arbusculus  Fisher,  represented  by 
a  female  and  nauplii  from  Hippasteria  californica  Fisher,  is  redescribed,  with  special  attention  to  the  female 
internal  anatomy.  Also,  two  new  species  are  described:  D.  fisheri  n.sp.,  represented  by  two  females,  from 
Pedicellaster  magister  megalabis  Fisher;  and  D.  punctata,  represented  by  females,  a  male,  metanauplii,  and 
ascothoracid  larvae,  from  Poraniopsis  inflata  (Fisher).  These  are  the  only  ascothoracids  so  far  described  from 
the  west  coast  of  the  Americas  south  of  Alaska. 


INTRODUCTION 

Ascothoracida  Lacaze-Duthiers,  1880,  is 
often  classified  (under  the  name  Ascothoracica 
Gruvel,  1905)  as  a  primitive  order  of  Cirripedia 
(e.g.,  Lacaze-Duthiers  1880,  1883;  Knipowitsch 
1892;  Gruvel  1905;  Newman  et  al.  1969),  but  ar- 
guments for  its  elevation  to  an  independent  sub- 
class have  also  been  made  (Wagin  1937,  1947, 
1976;  Grygier  1980b,  198 la).  The  taxon  now  com- 
prises about  40  species  in  10  genera,  all  parasites 
of  echinoderms  or  coelenterates  (reviewed  by 
Wagin  1976). 

Dendrogaster  Knipowitsch,  1890,  parasitic  in 
the  coelomic  cavity  of  sea-stars  (Asteroidea),  is 
the  largest  genus  in  the  subclass.  It  includes  over 
20  species  about  the  world,  the  taxonomy  and 
biology  of  which  have  been  studied  principally 
by  the  Russian  worker  Wagin  (especially  1950, 
1954,  1957;  summarized  1976).  Other  workers 
on  this  genus  have  included  Knipowitsch  (1890, 
1891,  1892),  Le  Roi  (1905,  1907),  Okada  (1925, 


1938,  1941),  Yosii  (1931),  Korschelt  (1933), 
Hickman  (1959),  Achituv  (1971),  Karande  and 
Oguro(1979,  1981a,  1 98 Ib),  and  Grygier  (198 la, 
1981b). 

Females  of  Dendrogaster  have  a  large, 
branched  mantle  (modified  carapace)  containing 
gut  diverticula,  gonads,  and  a  spacious  brood 
chamber.  The  main  body  is  reduced,  only  the 
first  antennae  and  mouth  parts  being  strongly 
developed.  Eggs  hatch  as  nauplii,  metanauplii, 
or  bivalved  ascothoracid  larvae  (Wagin  1948, 
1954).  Larvae  remain  within  the  brood  chamber 
until  the  ascothoracid  stage.  Males,  which  also 
live  in  the  brood  chamber,  have  an  ascothoracid 
larvalike  main  body  with  five  pairs  of  natatory 
appendages  and  a  muscular  abdomen  terminat- 
ing in  a  furca.  Their  carapace  lining  is  expanded 
posteriorly  into  a  pair  of  winglike  protrusions 
containing  testes  and  a  branch  of  the  gut  (Wagin 
1946,  1954). 

Dendrogaster  arbusculus  Fisher,  1911,  para- 


[443] 


444 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  18 


sitic  in  Hippasteria  californica  Fisher,  1905,  is 
the  only  named  ascothoracid  from  the  west 
coast  of  the  Americas  outside  Alaska,  although 
I  have  seen  a  specimen  of  Ascothorax  Djako- 
nov,  1914,  from  Monterey  Bay,  and  there  are 
two  undescribed  species  of  Dendrogaster  in  the 
Puget  Sound  region  (P.  Illg,  Univ.  of  Washing- 
ton, personal  communication).  Dendrogaster 
arbusculus  is  incompletely  known;  the  original 
description  (Fisher  1911)  consisted  only  of  the 
locality  of  collection,  the  host  sea-star,  and  a 
drawing  of  the  holotype  in  situ.  Yosii  (1931)  pre- 
sented a  schematic  diagram  of  the  mantle 
branching  pattern  based  on  Fisher's  drawing. 
This  species  was  included  in  the  genus  Myrio- 
cladus  Okada,  1925  (Okada  1925;  Yosii  1931; 
Kriiger  1940),  which  was  later  synonymized 
with  Dendrogaster  (Wagin  1950;  Hickman 
1959). 

Fisher  (191 1 ,  1928)  found  two  other  California 
sea-stars  infested  by  Dendrogaster,  Poraniopsis 
inflata  (Fisher),  1906,  and  Pedicellaster  magis- 
ter  megalabis  Fisher,  1928,  but  he  only  specified 
their  collection  localities.  His  tentative  identifi- 
cation of  these  parasites  as  D.  arbusculus  was 
never  confirmed  and,  as  shown  below,  was  mis- 
taken. 

METHODS  AND  MATERIALS 

All  of  Fisher's  finds  were  from  sea-stars  col- 
lected by  the  U.S.  Bureau  of  Fisheries  vessel 
ALBATROSS  off  California  in  1904;  station  data 
are  listed  below. 

Specimen  1,  holotype  of  Dendrogaster  arbus- 
culus; 5.4  km  SSE  of  Gull  Islet,  off  SE  coast  of 
Santa  Cruz  Island,  1220  m,  ALBATROSS  sta. 
4429,  14:IV:04;  host  species  Hippasteria  cal- 
ifornica (Fisher  1911:237,  404,  pi.  111). 

Specimen  2,  holotype  of  Dendrogaster  puncta- 
ta;  9.9  km  NW  of  Pt.  Pinos  Lighthouse,  Mon- 
terey Bay,  118-554  m,  ALBATROSS  sta.  4471, 
14:V:04;  host  species  Poraniopsis  inflata 
(Fisher  1911:264). 

Specimen  3 ,  paratype  fragments  of  Dendrogas- 
ter punctata;  same  station  as  holotype. 

Specimen  4,  holotype  of  Dendrogaster  fisheri; 
25.2  km  ssw  of  Pt.  Loma  Lighthouse,  San 
Diego,  940-960  m,  ALBATROSS  sta.  4334, 
14:IV:04;  host  species  Pedicellaster  magister 
megalabis  (Fisher  1928:66). 

Specimen  5,  paratype  of  Dendrogaster  fisheri; 
7.4  km  sw  of  Pt.  San  Pedro,  Santa  Cruz  Is- 


land, 817-932  m,  ALBATROSS  sta.  4427, 
9:111:04;  host  species  Pedicellaster  magister 
megalabis  (Fisher  1928:66). 

Dendrogaster-infested  sea-stars  or  isolated 
specimens  of  the  parasites  were  obtained  from 
the  California  Academy  of  Sciences  (CAS)  and 
the  National  Museum  of  Natural  History 
(USNM).  In  cases  where  a  sea-star  still  con- 
tained its  parasites,  the  host  was  dissected  and 
the  specimen  of  Dendrogaster  removed.  One 
previously  isolated  but  dried-out  mantle  frag- 
ment was  reconditioned  in  a  10%  trisodium 
phosphate  solution  overnight. 

At  least  one  female  of  each  species  was  dis- 
sected to  remove  appendages  and,  in  the  case 
of  the  holotype  of  D.  arbusculus,  the  internal 
organs.  Incisions  were  made  sparingly  in  each 
female  in  a  search  for  males  and  larvae.  A  male 
and  several  larvae  from  the  species  parasitic  in 
Poraniopsis  inflata,  as  well  as  several  larvae 
from  D.  arbusculus,  were  examined  whole  in 
lactic  acid;  some  larvae  of  each  species  were 
dissected.  Permanent  mounts  were  made  in  Tur- 
tox  CMC- 10  with  acid  fuchsin  or  in  glycerine 
jelly.  Drawings  were  done  with  the  aid  of  a  cam- 
era lucida. 

SYSTEMATIC  SECTION 

Ascothoracida  Lacaze-Duthiers,  1880 

Dendrogastridae  Gruvel,  1905 
Dendrogaster  Knipowitsch,  1890 

Dendrogaster  arbusculus  Fisher,  1911 

(Figures  1  &  2) 

Myriocladus  arbusculus:  OKADA  1925:371. 

MATERIAL. — One  female,  holotype,  deposited 
as  CAS  013 159. 

DIAGNOSIS. — Female  large.  Middle  piece  four 
times  as  long  as  main  branches;  small  "extra 
branch"  opposite  middle  piece.  Six  complexly 
ramifying  major  branches,  including  four  pos- 
terior secondary  branches  and  both  anterior  pri- 
mary branches.  Terminal  protuberances  conical, 
scattered.  Third  article  of  first  antenna  with  fu- 
sion seam,  proximodorsal  muscle,  and  dorsal 
seta.  Nauplii  about  1  mm  long,  with  setose  ap- 
pendages. Males  and  ascothoracid  larvae  un- 
known. 

DESCRIPTION  OF  FEMALE. — General  appear- 
ance. Fisher's  illustration  of  the  holotype  of  D. 
arbusculus  (1911:pl.  Ill,  fig.  1)  accurately  por- 
trays the  parasite  and  its  position  within  its  host. 
The  mantle  is  a  whitish  branched  sac  extending 


GRYGIER:  SEA-STAR  PARASITES 


445 


FIGURE  1.  Dendrogaster  arbusculus  Fisher;  holotype,  female.  («)  Schematic  diagram  of  mantle  branching  pattern,  terminal 
protuberances  omitted,  encircled  arrows  representing  broken  ends,  opposed  arrows  (lower  left)  indicating  reconstruction  (ori- 
entation of  distal  part  arbitrary),  (b)  Middle  piece,  dorsal  view,  (c)  First  antenna  with  musculature,  articles  numbered,  (d) 
Second  maxillae  with  musculature,  anterior  view,  (e)  Thoracic  part  of  nervous  system,  dorsal  view.  Explanation:  b,  primary 
branches;  b',  secondary  branches;  bn?,  buccal  nerve?;  c,  circumesophageal  connectives;  d,  distal  prong;  e,  "extra"  branch; 
m,  main  branches;  mp,  middle  piece;  s,  fusion  seam;  si,  distal  slit;  th,  thoracic  nerve  mass;  v,  ventral  prong.  Scale  bars  0.1 
mm,  except  1.0  mm  in  b. 


about  50  mm  in  the  preserved  state.  The  branch- 
es are  constricted  at  nodes  and  expanded  else- 
where, especially  distally.  The  very  thin  outer 
cuticle  of  the  mantle  covers  a  loose  network  of 
longitudinal  and  circular  muscle  fibers.  The  focal 
point  of  the  branching  pattern  (Fig.  la)  is  a  cla- 
vate  medial  protuberance  (middle  piece)  about 
10  mm  long  with  a  vertical  distal  slit  (Fig.  16). 


The  musculature  of  the  middle  piece  is  better 
developed  than  that  of  the  rest  of  the  mantle; 
prominent  circular  muscles  overlie  longitudinal 
muscular  sheets. 

Branching  pattern  (Fig.  la).  The  descriptive 
terminology  of  the  branching  pattern  is  modified 
from  that  of  Wagin  (1950,  1976).  A  pair  of  lat- 
erally directed  main  branches  arise  at  the  base 


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FIGURE  2.  Dendrogaster  arbusculus  Fisher;  nauplii.  (a)  Nauplius,  ventral  view,  diagrammatic,  (b)  First  antenna,  (c)  Second 
antenna,  (d)  Posterior  end  including  furcal  rudiments.  Explanation:  a,  first  antenna;  a',  second  antenna;  an,  anal  spine  or  seta; 
cr,  ctenate  ridges;  ds,  dorsal  shield;  end,  endopod;  ex,  exopod;  f,  furca:  md,  mandible;  sp,  furcal  spines;  y,  yolk.  Scale  bars 
0.1  mm. 


of  the  middle  piece,  each  about  2.5  mm  long. 
Opposite  the  middle  piece  is  a  short,  bifurcate 
"extra  branch."  The  main  branches  split  into 
anterior  and  posterior  primary  branches,  the  lat- 
ter quickly  dividing  into  a  pair  of  large  second- 
ary branches  on  each  side.  Further  ramifications 
of  these  six  major  branches  occur  by  an  irregular 
series  of  bifurcations  and  lateral  branchings, 
leading  finally  to  conical  terminal  protuberances 
sometimes  arranged  in  loose  trefoils. 

Internal  structure  of  mantle.  The  main  body, 
located  within  the  slit  at  the  distal  end  of  the 
middle  piece,  is  fused  to  the  mantle  dorsally  and 
laterally.  The  subchelate  first  antennae  and 
piercing-and-sucking  oral  cone  are  directed  ante- 
riorly. The  thorax  and  abdomen  are  almost  com- 
pletely reduced,  forming  a  sac  around  the  gut 
and  nervous  mass.  A  pair  of  large,  diaphanous 
gut  diverticula  pass  posteriorly  along  the  inside 
of  the  middle  piece,  extend  into  the  main 
branches,  and  follow  the  branching  of  the  mantle 


exactly.  The  ovaries,  attached  all  along  the  gut 
diverticula,  are  tissue  strands  or  tubules,  the  lat- 
eral follicles  of  which  each  contain  a  developing 
oocyte  (cf.  Okada  1941,  for  description  of  sim- 
ilar arrangement  in  another  species).  The  mantle 
cavity  is  a  brood  chamber  for  eggs  and  nauplii. 
First  antennae  (Fig.  Ic).  The  first  antennae  are 
four-segmented  and  subchelate,  0.7  mm  long. 
The  shape  of  the  first  article  could  not  be  deter- 
mined; its  musculature  consists  of  two  dorsal 
flexors  and  two  ventral  extensors  of  the  second 
article  and  a  long  flexor  inserted  on  the  proxi- 
moventral  corner  of  the  third  article.  The  second 
article  is  rectangular.  A  flexor  and  an  extensor 
muscle  run  from  its  proximodorsal  corner  to  the 
proximal  corners  of  the  third  article;  another 
flexor  runs  from  its  proximoventral  side  to  the 
proximoventral  corner  of  the  third  article.  The 
third  article  is  subrectangular,  with  its  distal 
edge  facing  somewhat  dorsally.  A  seam  indicat- 
ing the  fusion  of  two  articles  cuts  across  the 


GRYGIER:  SEA-STAR  PARASITES 


447 


proximodorsal  corner;  there  is  a  seta  near  the 
distal  end  of  the  seam  and  a  muscle  within  the 
corner  set  off  by  it.  The  fourth  article  is  oval, 
fitting  lengthwise  into  the  distal  end  of  the  third. 
There  are  two  anterior  processes,  the  first  a 
small  mound  with  two  setae,  the  other  cylindri- 
cal with  two  setae  on  the  right  first  antenna, 
three  on  the  left.  There  is  a  seta  at  the  base  of 
the  heavy  distal  claw.  Two  claw-retracting  mus- 
cles run  from  a  thickening  anterior  to  the  claw 
(cf.  Grygier  1981b);  one  attaches  to  the  side 
of  the  article,  the  other  to  its  basal  apodeme. 
This  apodeme  extends  far  into  the  third  article; 
a  very  powerful  fan  of  flexor  muscles  reaches 
from  it  to  the  ventral  and  proximal  sides  of  that 
article.  A  less  powerful  extensor  muscle  runs 
alongside  the  flexors  from  the  ventral  base  of 
the  fourth  article  to  the  proximal  end  of  the 
third. 

Oral  cone.  The  oral  cone  is  directed  anterior- 
ly. The  labrum  forming  the  sheath  of  the  cone 
is  drawn  out  laterally  and  ventrally,  almost  en- 
closing the  mouth  parts.  The  basal  part  is  swol- 
len with  the  insertions  of  the  buccal  pump 
muscles.  The  only  mouth  parts  are  the  harpoon- 
shaped  second  maxillae  (Fig.  Id),  which  pro- 
trude from  the  cone.  They  are  fused  for  half  their 
length.  The  distal  prong  at  the  tip  of  each  con- 
sists of  a  thick  point  and  a  membranous  flange. 
The  ventral  hook  is  bent  laterally  downwards 
and  is  more  heavily  chitinized  than  the  distal 
prong.  A  muscle  runs  from  its  base  to  the  base 
of  the  mouth  part,  where  it  ends  at  a  transverse 
wrinkle  indicating  an  articulation.  There  are 
many  short  muscles  more  proximally. 

Internal  anatomy.  The  buccal  pump  is  a  mass 
of  circular,  longitudinal,  and  radial  muscles  oc- 
cupying the  bulk  of  the  oral  cone.  It  encloses 
the  chitin-lined  esophagus  which  leads  into  the 
midgut,  from  which  the  digestive  diverticula 
emerge.  In  other  species  of  Dendrogaster,  the 
gut  is  blind  (Knipowitsch  1892;  Le  Roi  1907; 
Okada  1925;  Wagin  1954);  it  was  not  possible  to 
confirm  that  this  is  the  case  here. 

The  nervous  system  (Fig.  le)  is  highly  con- 
centrated in  the  thorax.  A  pair  of  round,  medi- 
ally connected  cerebral  ganglia  lie  anterior  to  the 
esophagus;  though  they  are  said  to  give  off  large 
antennular  nerves  (Wagin  1954),  these  were  not 
found.  Lateral  connectives  from  the  cerebral 
ganglia  pass  around  the  esophagus,  giving  off  a 
pair  of  bifurcate  buccal  (?)  nerves.  The  connec- 


tives are  linked  by  a  commissure  before  they 
enter  the  anteroventral  part  of  the  thoracic  nerve 
mass.  From  this  mass,  which  is  not  divided  into 
ganglia,  a  pair  of  very  large  nerves  arise  laterally 
to  innervate  the  mantle.  Several  small  nerves 
arise  on  each  side  near  their  bases,  two  or  three 
anteriorly  and  one  posterodorsally.  Two  pairs  of 
fairly  large  nerves  arise  laterally  from  the  pos- 
terior end  of  the  thoracic  mass,  one  dorsal  and 
one  ventral.  It  is  impossible  to  say  what  struc- 
tures all  these  nerves  innervate;  no  such  prolif- 
eration of  nerves  has  been  described  in  Dendro- 
gaster previously  (cf.  Wagin  1954). 

A  pair  of  large,  wrinkled  saclike  organs  were 
removed  with  the  second  maxillae.  These  may 
be  the  maxillary  glands,  though  no  ducts  could 
be  followed  nor  openings  on  the  maxillae  locat- 
ed. 

A  cavity  within  the  main  body  contained  a 
white,  friable  substance  which  resembled  yolk 
except  for  its  color.  A  similar  organ  has  been 
described  in  a  new  Antarctic  species  of  Dendro- 
gaster, where  it  was  considered  a  vitellogenous 
organ  (Grygier  1981b;  also  see  Grygier  1980a). 
It  more  likely  represents  the  proximal  part  of 
the  oviduct. 

GAMETES  AND  LARVAE. — Eggs.  Hundreds  of 
eggs  were  found  within  the  mantle  cavity.  They 
are  yellow  and  subspherical,  averaging  0.51 
mm  x  0.45  mm. 

Sperm.  Patches  of  sperm  cells  are  attached  to 
membranes  within  the  mantle.  Their  heads  are 
about  4.2  fj,m  long,  1.7-2.0  /xm  wide,  tapered 
slightly  anteriorly,  with  a  small  terminal  aero- 
some.  The  midpiece  is  granular  and  tapers  pos- 
teriorly for  about  6.4  /im.  The  tail  is  at  least  20.4 
/im  long  and  probably  much  longer.  These 
sperm  are  similar  to  those  of  other  species  of 
Dendrogaster  studied  by  light  microscopy  (Le 
Roi  1907;  Okada  1941)  and  by  electron  micros- 
copy (Grygier  198 la).  If  these  sperm  are  not  au- 
tochthonous but  are  actually  ingested  starfish 
sperm,  they  should  have  round  heads  (Dan 
1968),  not  the  observed  oblong  heads. 

MALES. — It  is  puzzling  that  no  males  were 
found  in  this  female.  Nauplii  occupied  the  brood 
chamber;  therefore,  fertilization  must  have  tak- 
en place.  Parthenogenesis  has  been  suggested  in 
D.  murmanensis  Wagin,  1950  (Kluge,  unpub- 
lished data  cited  by  Wagin  1947),  and  D.  reper- 
tus  Le  Roi,  nom.  nud.  (Le  Roi,  unpublished  data 
cited  by  Kriiger  1920),  but  since  sperm  were 


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present,  it  is  unlikely  to  have  been  the  case  here. 
One  possible  case  of  hermaphroditism  is  report- 
ed. Okada  (1941)  found  that  the  anterior  region 
of  the  ovary  in  D.  okadai  (Yosii,  1931)  produced 
spermlike  cells  rather  than  ova.  He  did  not  ex- 
plain why  he  did  not  consider  these  true  sperm. 
No  obvious  testes  were  found  in  the  present 
specimen,  but  internal  investigations  were  min- 
imized to  preserve  the  integrity  of  the  holotype. 
It  is  also  possible  that  a  dwarf  male  was  lost 
when  the  mantle  was  damaged  upon  removal 
from  the  host. 

NAUPLII  (Fig.  2). — About  24  nauplii,  each 
about  1.0  mm  long  and  0.9  mm  wide,  were  found 
(Fig.  2a).  The  dorsal  shield  is  bowl-shaped,  oval 
in  dorsal  view,  with  a  broad  rim.  The  main  body 
of  the  nauplius  completely  fills  the  bowl's  con- 
cavity. It  has  three  pairs  of  naupliar  appendages, 
a  simple  labrum,  and  a  rudimentary  furca,  but 
no  nauplius  eye.  The  first  antennae  are  unira- 
mous  and  unsegmented,  and  have  three  long  and 
two  short  distal  setae  (Fig.  2b).  The  second  an- 
tennae and  mandibles  are  alike  (Fig.  2c),  with 
an  obscurely  divided  protopod,  an  exopod 
formed  of  about  13  annuli,  the  last  6  or  so  each 
bearing  a  long  seta,  and  an  endopod  half  as  long 
as  the  exopod  with  about  four  indistinct  articles 
and  three  setae.  The  labrum  is  small,  its  poste- 
rior margin  acute.  The  furca  (Fig.  2d)  consists 
of  a  pair  of  conical  mounds,  each  with  a  spine 
at  the  apex.  A  long,  flexible  anal  spine  arises 
between  and  dorsal  to  them.  Between  the  la- 
brum and  furca,  more  pronounced  along  the 
midline  and  posteriorly,  are  a  number  of  ctenate 
cuticular  ridges. 

AFFINITIES. — The  elucidation  of  interspecific 
relationships  among  the  species  of  Dendrogas- 
ter  has  so  far  proved  intractable.  Okada' s  (1925) 
segregation  of  several  species  (including  D.  ar- 
busculus)  into  a  new  genus,  Myriocladus,  is  no 
longer  accepted.  Provisional  attempts  to  corre- 
late morphology  of  the  parasites  with  the  phy- 
logeny  of  their  host  sea-stars  have  been  unsat- 
isfactory (Yosii  1931;  Grygier  1981b). 

Wagin  (1976)  proposed  another  approach 
based  on  his  investigations  (Wagin  1947,  1948, 
1954)  of  embryonic  and  larval  development  in 
ascothoracids,  including  species  of  Dendrogas- 
ter.  He  distinguished  three  kinds  of  larval  de- 
velopment in  this  genus.  Dendrogaster  astro- 
pectinis  (Yosii,  1931)  and  D.  beringensis  Wagin, 
1957,  hatch  from  the  eggshell  as  nauplii.  Other 


species  hatch  as  metanauplii,  and  yet  others  as 
ascothoracid  larvae.  Wagin  considered  the  first 
variation  the  most  primitive  and  suggested  that 
the  deep  bathyal  habitat  (2000-4000  m)  of  the 
two  species  exhibiting  it  was  the  critical  factor 
in  its  retention.  Dendrogaster  arbusculus  hatch- 
es as  a  nauplius  that  is  more  perfectly  formed 
than  that  of  D.  astropectinis,  and  the  present 
specimen  was  captured  at  shallow  bathyal 
depths  (1200  m).  These  facts  may  not  indicate 
affinities  between  D.  arbusculus  and  either  D. 
astropectinis  or  D.  beringensis,  however,  be- 
cause the  presence  of  a  free  naupliar  stage  is  a 
plesiomorphic  feature. 

Neither  of  the  other  two  species  remotely  re- 
sembles D.  arbusculus  in  its  branching  patterns. 
Like  D.  arbusculus,  females  of  D.  astropectinis 
have  a  fusion  seam  and  a  proximodorsal  muscle 
in  the  third  article  of  the  first  antenna.  This  con- 
formation, which  is  also  plesiomorphic,  is  not 
limited  to  these  two  species,  however,  being 
found  also  in  D.  iwanowi  Wagin,  1950,  D.  tas- 
maniensis  Hickman,  1959,  and  both  new  species 
described  below.  The  occurrence  of  a  single  seta 
on  this  article  is  limited  to  D.  arbusculus  and 
one  of  the  new  species  below.  Dendrogaster 
ludwigi  Le  Roi,  1905,  and  D.  dogieli  Wagin, 
1950,  the  only  other  species  in  the  genus  with  an 
"extra  branch"  like  D.  arbusculus,  otherwise 
have  unique  branching  patterns  not  resembling 
that  of  the  present  species. 

Dendrogaster  fisheri,  new  species 

(Figure  3) 

Dendrogaster  cf.  arbusculus:  FISHER  1928:66. 

MATERIAL. — Two  females.  Holotype  from 
ALBATROSS  sta.  4334  deposited  as  USNM 
184574;  paratype  from  ALBATROSS  sta.  4427  as 
USNM  184575.  Holotype  occupied  four  arms  of 
host,  middle  piece  directed  outward  in  interra- 
dius  (Fig.  3« );  paratype  occupied  only  two  arms 
of  host. 

DIAGNOSIS. — Females  small  to  medium-sized, 
with  four  equally  developed  primary  branches. 
Short  middle  piece  more  than  twice  as  long  as 
main  branches;  short,  lobate  secondary  branch- 
es arising  alternately  from  primary  branches. 
Third  article  of  first  antenna  with  fusion  seam, 
proximodorsal  muscle,  and  two  or  three  dorsal 
setae,  one  removed  from  rest.  Males,  nauplii, 
and  ascothoracid  larvae  unknown. 


GRYGIER:  SEA-STAR  PARASITES 


449 


FIGURE  3.  Dendrogasterfisheri,  new  species;  female,  (a)  Holotype  in  situ  in  host  sea-star  Pedicellaster  magister  megalabis 
Fisher,  dorsal  view,  (b)  Holotype,  ventral  view,  (c)  Paratype,  ventral  view,  (d)  First  antenna  of  paratype  with  musculature, 
articles  numbered,  (e)  First  antenna  of  holotype,  distal  articles  (numbered).  (/)  Tip  of  second  maxilla,  lateral  view.  Explanation: 
b,  primary  branches;  b',  secondary  branches;  eg,  eggs;  mp,  middle  piece;  s,  fusion  seam. 


DESCRIPTION. — General  appearance.  Den- 
drogasterfisheri is  of  small  to  medium  size,  the 
greatest  extent  of  the  branches  being  26  mm  in 
the  holotype  (Fig.  3a,  b)  and  14  mm  in  the  para- 
type (Fig.  3c).  The  mantle  surface  is  smooth 
with  clearly  visible  longitudinal  muscles.  Its  col- 
or in  the  preserved  state  is  pale  brown  to  pale 
yellow.  The  oral  cone  and  left  first  antenna  of 
the  holotype  protruded  through  the  distal  slit  of 
the  middle  piece  before  dissection,  but  no  part 
of  the  paratype's  main  body  was  visible  exter- 
nally. 

Branching  pattern.  The  middle  piece  of  the 
holotype  is  2  mm  long,  bluntly  conical  with  a 
distal  slit.  The  main  branches  are  very  short 
(less  than  1  mm),  each  dividing  into  a  pair  of 
primary  branches  9-12  mm  long  in  the  holotype, 
6-9  mm  long  in  the  paratype.  The  anterior  and 
posterior  primary  branches  are  equally  devel- 


oped. Many  short  secondary  branches  arise  al- 
ternately from  the  anterior  and  posterior  sides 
of  the  primary  branches,  the  proximal  ones 
slightly  larger  than  the  distal  ones,  each  bearing 
several  blunt  terminal  protuberances. 

First  antennae  (Fig.  3d,  e).  The  four- seg- 
mented, subchelate  first  antennae  are  massive, 
0.5  mm  long  in  the  paratype  (Fig.  3d),  larger  in 
the  holotype  (Fig.  3e).  The  basal  article  is  tri- 
angular, attached  to  the  head  by  its  longest  side, 
and  with  its  sharpest  apex  uppermost;  it  con- 
tains the  flexor  and  extensor  muscles  of  the  sec- 
ond article  and  a  flexor  muscle  of  the  third  ar- 
ticle. The  second  article  is  rectangular,  smaller 
and  shorter  than  the  first,  and  contains  flexor 
and  extensor  muscles  of  the  third  article.  The 
third  article  is  an  irregular  quadrilateral  higher 
than  long,  with  a  fusion  seam  across  its  proxi- 
modorsal  corner  and  two  or  three  dorsal  setae 


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distal  to  the  seam  (one  isolated  far  distally);  it 
contains  flexor  and  extensor  muscles  of  the 
fourth  article  and  an  isolated  muscle  in  the  cor- 
ner delimited  by  the  fusion  seam.  The  fourth, 
distalmost  article  is  short,  almost  square,  with 
a  distal  claw  and  two  claw  retractor  muscles.  It 
is  armed  with  a  small  seta  lateral  to  the  claw,  a 
process  on  the  distal  face  with  three  setae,  and 
a  small  seta  at  the  base  of  the  claw. 

Oral  cone.  The  only  obvious  mouth  parts,  the 
second  maxillae,  are  enclosed  in  a  sheathlike 
oral  cone  formed  from  the  labrum.  The  tips  of 
the  second  maxillae  are  bifurcate  (Fig.  3/);  the 
ventral  hook  can  be  bent  laterally  and  poste- 
riorly with  respect  to  the  distal  prong.  The  tips 
are  directed  away  from  each  other  when  the 
mouth  parts  protrude  from  the  oral  cone. 

GAMETES  AND  LARVAE. — In  both  specimens 
a  large  number  of  eggs  about  0.5  mm  in  diameter 
distend  the  primary  branches  and  bend  the  sec- 
ondary branches  ventrally  (Fig.  3b).  No  dwarf 
males  or  larvae  were  found  in  a  cursory  search 
of  both  specimens. 

AFFINITIES. — Fisher  (1928)  tentatively  as- 
signed these  specimens  to  D.  arbusculus .  There 
is  no  real  similarity,  however,  since  D.  fisheri 
has  only  four  major  branches  with  no  compli- 
cated higher-order  branching,  a  relatively  short- 
er middle  piece  than  D.  arbusculus,  and  no 
"extra  branch"  opposite  the  middle  piece.  At 
the  time  of  Fisher's  publication,  only  four  other 
species  of  Dendrogaster  had  been  named  and 
the  extent  of  intraspecific  variation  of  branching 
patterns,  still  incompletely  known  (but  see  Ka- 
rande  and  Oguro  198  Ib),  could  not  be  estimated. 
Since  the  two  available  specimens  of  D.  fisheri 
are  alike  in  most  respects,  including  their  com- 
mon host,  I  am  proposing  a  new  species  for 
them.  In  recognition  of  his  having  informed  zo- 
ologists of  the  existence  of  these  specimens,  I 
take  pleasure  in  naming  the  species  after  W.  K. 
Fisher. 

Dendrogaster  fisheri  superficially  resembles 
young  specimens  of  D.  arctica  Korschelt,  1933, 
which  also  have  four  equal  primary  branches 
with  lobular  side  branches  (Wagin  1950),  but  in 
D.  arctica  the  secondary  branches  are  simple, 
lacking  terminal  protuberances.  The  first  anten- 
na has  no  fusion  seam  or  associated  muscle, 
only  two  setae  on  the  third  article,  and  a  single 
seta  proximal  to  the  process  on  the  fourth  article 
(Wagin  1950).  Dendrogaster  arctica  inhabits  the 


Bering  Sea  (Fisher  1930;  Wagin  1950)  and  Hud- 
son Bay  (undocumented  record  on  distribution 
map  by  Wagin  1970),  and  parasitizes  the  sea-star 
Leptasterias  groenlandica  (Lu'tken),  1857. 

Dendrogaster  ludwigi  also  has  four  primary 
branches.  The  secondary  branches,  though 
short,  are  so  densely  covered  with  terminal 
protuberances  that  the  primary  branches  are 
completely  obscured,  whereas  in  D.  fisheri  the 
primary  branches  are  easily  discernible.  Den- 
drogaster ludwigi  lives  in  shallow  water  from 
Japan  to  Australia  (Le  Roi  1905,  1907;  Yosii 
1931;  Kenny  1959);  its  range  is  unlikely  to  ex- 
tend to  the  east  Pacific,  especially  in  waters  as 
deep  as  D.  fisheries  habitat  (800-1000  m). 

In  no  other  known  species  of  Dendrogaster 
is  the  mantle  constructed  around  four  relatively 
simple  primary  branches. 

Dendrogaster  punctata,  new  species 

(Figure  4) 

Dendrogaster  cf.  arbusculus:  FISHER  1911:264. 

MATERIAL. — One  female,  holotype,  deposited 
as  USNM  184573;  several  mantle  fragments 
from  same  host,  paratype  lot,  deposited  as  CAS 
018895. 

DIAGNOSIS. — Females  medium-sized  with 
complexly  branched  mantle.  Short,  cylindrical 
middle  piece  little  longer  than  main  branches. 
Primary  and  secondary  branches  dichotomous, 
better  developed  posteriorly;  more  distal 
branches  either  unbranched  and  elongate  or  var- 
iously branched,  ending  in  trefoil- shaped  clus- 
ters of  terminal  protuberances.  Mantle  exten- 
sively pitted.  Third  article  of  first  antenna  with 
weak  fusion  seam,  proximodorsal  muscle,  and 
dorsal  seta.  Males  with  elongate,  cylindrical 
posterior  protrusions;  testes  ribbonlike.  Third 
article  of  first  antenna  in  males  and  ascothoracid 
larvae  without  proximodorsal  muscle,  but  with 
two  long,  bifid  setae.  Anterior  face  of  fourth  ar- 
ticle with  straplike  aesthetasc,  seta,  and  process 
with  three  setae.  Late  metanauplii  lacking  an- 
tennae. 

DESCRIPTION  OF  FEMALE. — General  appear- 
ance (Fig.  4a).  The  mantle  extends  26  mm  when 
the  branches  are  spread  out,  but  in  situ  the 
branches  are  curled  and  intertwined  (fixation 
artifact?).  The  mantle  is  beige  and  fairly  tough 
in  the  preserved  state.  Its  surface  is  uneven,  pit- 
ted by  irregular  small  depressions  (Fig.  4b).  [The 


GRYGIER   SEAOBSTAR  PARASITES 


451 


FIGURE  4.  Dendrogaster  punctatus,  new  species,  (a)  Holotype,  female,  overall  view,  (b)  Detail  of  same  showing  depressions 
in  mantle  (stippled),  (c)  First  antenna  of  same  with  musculature,  articles  numbered,  (d)  Male,  dorsal  view,  (e)  Male,  lateral 
view,  left  valve  removed,  natatory  setae  omitted  on  thoracopods  and  furca.  (/)  Male,  second  thoracopod  with  natatory  setae. 
(g)  Late  metanauplius,  ventral  view,  (h)  Carapace  valve  of  ascothoracid  larva,  interior  view,  anterior  end  right,  (i)  Main  body 
of  ascothoracid  larva,  most  thoracopod  solution  omitted,  first  antenna!  articles  numbered.  Explanation:  a,  first  antenna;  a'?, 
second  antenna?;  ad,  adductor  muscle;  aes,  aesthetasc;  b,  primary  branches;  b',  secondary  branches;  ds,  dorsal  shield;  ep, 
epaulet;  es,  esophagus;  f,  furcal  rami;  g,  guard  spinules;  gu,  midgut  (diverticula  indicated  by  dashed  lines  in  </):  1,  labrum;  m, 
main  branch;  mp,  middle  piece;  mx,  second  maxilla;  oc,  oral  cone;  p,  penis;  pr,  posterior  protrusion;  s,  fusion  seam;  t,  testes; 
tr,  thoracopod  rudiments;  v,  carapace  valve.  Scale  bars  1.0  mm  in  a,  b,  d;  otherwise  0.1  mm. 


452 


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specific  name  is  derived  from  this  sculpturing 
(Latin  punctatus:  prick  or  point).] 

Branching  pattern.  The  middle  piece  is  rather 
thick  and  3.5  mm  long,  slightly  longer  than  the 
main  branches.  The  latter  bifurcate  into  short 
primary  branches,  each  of  which  bifurcates 
again  into  secondary  branches,  the  posterior 
ones  ramifying  a  little  more  complexly  than  the 
anterior  ones.  Some  secondary  branches  bifur- 
cate or  trifurcate;  others  are  palmate.  Some  ter- 
tiary branches  are  simple  elongate  lobes;  usu- 
ally, though,  higher-order  branches  end  in 
terminal  protuberances  arranged  in  trefoils. 

First  antennae  (Fig.  4c).  The  first  antennae  are 
four-segmented  and  subchelate.  The  dorsal  part 
of  the  triangular  first  article  overlaps  part  of  the 
tall,  rectangular  second  article.  The  third  article 
is  a  rounded  quadrilateral  with  a  faint  fold  in- 
dicating a  fusion  seam,  a  muscle  in  the  corner 
so  delimited,  and  one  large  dorsal  seta.  The 
fourth  article  is  rhombic  with  a  strong  distal 
claw.  Its  anterior  face  is  armed,  progressively 
more  distally,  with  two  setae,  a  process  with 
two  or  three  terminal  setae,  and  a  smaller  seta 
arising  from  the  side  of  the  article.  There  is  a 
small  seta  near  the  base  of  the  claw.  The  mus- 
culature is  similar  to  that  described  in  D.  ar- 
busculus  and  D.  fisheri. 

Oral  cone.  The  sheath  of  the  oral  cone  is 
formed  from  the  expanded  lab  rum.  The  mouth 
parts  were  not  examined. 

DESCRIPTION  OF  FEMALE  FRAGMENTS. — It  is 
not  possible  to  determine  how  the  several  man- 
tle fragments  found  together  with  the  holotype 
fit  together  to  form  one  or  more  complete  spec- 
imens, especially  since  no  middle  pieces  were 
among  them.  Two  small  fragments  may  be  from 
a  single  specimen  similar  to  the  holotype.  Two 
others  may  be  from  a  larger  specimen  with  fewer 
terminal  protuberances  in  trefoils  than  the  ho- 
lotype and  a  greater  proportion  of  unbranched, 
elongate  higher-order  branches.  All  fragments 
had  pitted  surfaces  like  the  holotype. 

DESCRIPTION  OF  MALE. — General  appearance 
(Fig.  4d,  e).  The  single  dwarf  male  found  in  one 
of  the  female  mantle  fragments  is  bivalved  with 
a  cylindrical  protrusion  arising  posteriorly  from 
each  valve.  The  valves  are  about  0.63  mm  long 
and  are  held  open  for  a  total  width  of  0.70  mm. 
One  posterior  protrusion  is  1.6  mm  long,  the 
other  1.4  mm.  A  midgut  diverticulum  partially 
enveloped  by  longitudinal  strips  of  testicular  tis- 


sue reaches  nearly  to  the  end  of  each  protrusion 
(Fig.  4d).  The  main  body,  suspended  beneath 
the  valves,  has  four-segmented,  subchelate  first 
antennae,  an  oral  cone,  five  thoracomeres  with 
natatory  limbs,  and  a  five-segmented  abdomen 
terminating  in  a  pair  of  furcal  rami  (Fig.  4e). 

First  antennae  (Fig.  4e).  The  first  antennae  are 
massive  pincers  protruding  from  the  carapace. 
The  first,  triangular  article  is  the  largest;  the  sec- 
ond and  third  are  rectangular,  the  latter  with  a 
proximodorsal  fusion  seam  and  a  pair  of  large 
terminally  bifid  setae.  The  fourth  article  is  small- 
er than  the  others  and  bears  a  distal  claw  op- 
posing the  two  setae  on  the  third  article.  A  strap- 
like  sensory  aesthetasc  almost  twice  as  long  as 
the  first  antenna  arises  from  the  ventroproximal 
corner.  Distal  to  this  on  the  anterior  face  is  a 
large  seta  and,  farther  distally,  a  fixed  process 
bearing  three  setae.  Two  long  setae  flank  the 
claw,  and  a  third  arises  at  its  base.  All  these 
setae  narrow  abruptly  to  a  sharp  point.  Little 
can  be  made  of  the  musculature,  but  there  is  no 
muscle  in  the  part  of  the  third  article  delimited 
by  the  fusion  seam. 

Oral  cone  (Fig.  4e).  The  oral  cone  is  narrow. 
The  labral  margins  do  not  meet  posteriorly.  The 
distal  and  posterior  prongs  of  the  second  max- 
illae, the  only  mouth  parts,  are  strongly  chitin- 
ized. 

Thorax.  The  oral  cone  and  first  pair  of  tho- 
racopods  are  widely  separated  (Fig.  4e}.  A  pair 
of  ventrolateral  flanges  extend  from  the  thora- 
copods  to  the  first  antennae,  partly  overlapping 
the  oral  cone.  The  last  thoracomere  has  a  small 
posterolateral  epaulet  (Fig.  4e).  The  thoraco- 
pods  consist  of  a  coxa,  a  basis,  a  Inarticulate 
endopod  (biarticulate  in  the  fifth  pair),  and  a  bi- 
articulate  exopod  (Fig.  4/).  The  coxae  of  the  first 
two  pairs  each  bear  a  large  lateral  seta;  the  first 
four  pairs  have  a  medial  seta  on  the  basis  and 
their  endopods  have  one  seta  on  the  second  ar- 
ticle and  three  terminal  setae.  All  the  exopods 
terminate  in  three  setae. 

Abdomen.  There  is  a  short,  distinctly  bilobed 
penis  rudiment  on  the  venter  of  the  first  abdom- 
inal segment  (Fig.  4e).  The  furcal  rami  are  al- 
most square  with  four  long  medial  setae,  three 
arising  basally,  the  fourth  distally  near  the  dorsal 
margin.  Of  the  four  terminal  setae,  the  most  ven- 
tral is  less  than  half  the  length  of  the  others. 

DESCRIPTION  OF  LARVAE. — Metanauplii  (Fig. 
4g).  All  the  mantle  fragments  contained  numer- 


GRYGIER:  SEA-STAR  PARASITES 


453 


ous  late  metanauplii  about  0.55  mm  long  and 
0.35  mm  wide.  The  dorsal  shield  is  folded  ven- 
trally  at  the  sides  and  has  a  pronounced  anterior 
indentation,  presaging  the  bivalved  carapace  of 
the  ascothoracid  larva.  The  first  antennal  rudi- 
ments are  cylindrical  with  several  short  terminal 
setae.  Eyes  and  second  antennae  are  lacking.  A 
pair  of  stylets  (second  maxillae?)  protrude  from 
beneath  the  large  labrum.  No  other  mouth  parts 
are  visible.  There  are  five  pairs  of  stubby  tho- 
racopod  rudiments  bearing  setae,  as  well  as  a 
pair  of  unerupted  furcal  rami  with  numerous 
short  distal  setae. 

Ascothoracid  larvae  (Fig.  4/z,  /).  These  larvae 
are  very  similar  to  the  male  except  that  they  lack 
posterior  protrusions.  The  carapace  valves  (Fig. 
4/z)  are  about  0.60  mm  long  and  0.35  mm  high. 
The  posteroventral  margin  of  the  inner  cuticle 
is  lined  with  two  rows  of  guard  spinules.  On  the 
main  body  (Fig.  4/),  small  lobes  posterior  to  the 
first  antennae  may  be  vestiges  of  second  anten- 
nae (Okada  1938;  Hickman  1959).  The  oral  cone 
and  thoracopods  are  not  as  far  removed  from 
each  other  as  they  are  in  the  male,  but  a  space 
remains  corresponding  to  the  lost  first  pair  of 
thoracopods.  The  coxae  of  the  first  two  pairs  of 
developed  thoracopods  are  armed  externally 
with  a  fringe  of  fine  hairs  and  a  seta.  Other 
details  of  setation  are  unclear,  but  are  probably 
similar  to  those  in  the  male.  The  main  bodies  of 
the  ascothoracid  larvae  and  males  are  alike  in 
all  other  respects. 

AFFINITIES. — Fisher  (1911)  suggested  that 
these  specimens  belonged  to  D.  arbusculus.  The 
complexity  of  the  branching  pattern  is  compa- 
rable, but  numerous  differences  argue  against 
this  identification.  The  mantle  of  the  holotype 
of  D.  arbusculus  is  much  more  delicate  than 
that  of  D.  punctata  and  lacks  depressions.  The 
middle  piece  is  relatively  much  shorter  in  D. 
punctata,  which  also  lacks  an  "extra  branch" 
opposite  the  middle  piece.  The  terminal  protu- 
berances are  well  expressed  and  rounded  in  D. 
punctata,  but  are  less  pronounced  in  the  other 
species.  The  third  article  of  the  first  antenna  is 
identically  armed  in  both  species,  but  the  two 
proximal  setae  on  the  fourth  article  are  on  a 
mound  in  D.  arbusculus,  not  in  D.  punctata. 
The  metanauplii  and  ascothoracid  larvae  of  D. 
punctata  are  only  about  half  the  size  of  the  nau- 
plii  of  D.  arbusculus. 

Four  other  species  of  Dendrogaster  have  ter- 


minal protuberances  arranged  in  trefoils.  Den- 
drogaster rimskykorsakowi  Wagin,  1950,  is  not 
complexly  branched  and  the  clusters  of  terminal 
protuberances  are  widely  spaced  (Wagin  1950). 
Dendrogaster  okadai  has  oppositely  pinnate 
higher-order  branches  (Yosii  1931),  and  D.  lud- 
wigi  does  not  branch  beyond  the  tertiary  order. 
The  new  Antarctic  species  agrees  with  D.  punc- 
tata in  having  several  dichotomies  followed  by 
trichotomies  in  the  higher  branching  orders,  but 
it  is  much  more  robust  than  D.  punctata  and  has 
very  distinctive  first  antennae  (Grygier  1980a, 
1981b). 

The  metanauplii  of  D.  punctata  are  peculiar 
in  lacking  rudiments  of  the  second  antennae  and 
the  first  pair  of  thoracopods  (possibly  also  lack- 
ing mandibles  and  first  maxillae,  but  these 
mouth  parts  may  be  hidden  under  the  labrum). 
Wagin  (1954,  1976)  reported  the  presence,  albeit 
in  reduced  form,  of  all  these  appendages  through 
the  metanaupliar  stages  in  several  other  species 
of  Dendrogaster,  although  the  first  thoracopods 
were  always  lost  in  the  molt  to  the  ascothoracid 
larva.  He  may  have  been  conservative  in  de- 
scribing only  three  ontogenetic  sequences 
among  the  species  of  this  genus. 

Among  ascothoracid  larvae  and  males,  the 
long  bifid  setae  on  the  third  article  of  the  first 
antennae  are  unique  to  D.  punctata.  In  sum,  this 
species  seems  to  have  no  known  close  relatives 
elsewhere  in  the  genus. 

ACKNOWLEDGMENTS 

I  extend  thanks  to  Mr.  Dustin  D.  Chivers 
(CAS)  and  Dr.  David  Pawson  (USNM)  for  lo- 
cating the  specimens  of  Dendrogaster  discussed 
in  this  paper,  Dr.  Paul  Illg  (University  of  Wash- 
ington) for  enlightening  personal  discussions  of 
his  own  work  on  this  genus,  and  Dr.  William  A. 
Newman  (Scripps  Institution  of  Oceanography) 
for  allotting  me  space  in  his  laboratory  and  crit- 
icizing earlier  drafts  of  this  paper.  This  work,  a 
Contribution  of  Scripps  Institution  of  Oceanog- 
raphy, new  series,  was  done  during  the  tenure 
of  an  NSF  Graduate  Fellowship  and  was  partly 
supported  by  NSF  Grant  DEB78- 15052. 

LITERATURE  CITED 

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5(3):249-263. 

WAGIN,  V.  L.  [SPELLED  VAGIN].  1937.  Die  Stellung  der  As- 
cothoracida ord.  nov.  (Cirripedia  Ascothoracica  Gruvel 
1905)  im  System  der  Entomostraca.  C.  R.  (Dokl.)  Acad. 
Sci.  URSS  15(5):273-278. 

[SPELLED  VAGHIN].  1946.  On  males  of  Dendrogaster- 

idae  (Ascothoracida,  Entomostraca).  C.  R.  (Dokl.)  Acad. 
Sci.  URSS  52(3):273-276. 

.   1947.  Ascothorax  ophioctenis   and  the  position  of 


Ascothoracida  Wagin  in  the  system  of  the  Entomostraca. 
Acta  Zool.  (Stockh.)  27  [dated  1946]:  155-267. 
.    1948.    [On  types  of  larval  development  in  Dendro- 


gasteridae  (Ascothoracida,  Entomostraca).]  [In  Russian.] 
Dokl.  Akad.  Nauk  SSSR  59(2):387-390. 

-.    1950.    [On  new  parasitic  crustaceans  of  the  family 


Dendrogasteridae  (order  Ascothoracida).]  [In  Russian.]  Tr. 
Leningr.  O-va.  Estestvoispyt.  70(4):3-89. 
.    1954.    [On  structure,  larval  development,  and  meta- 


morphosis in  Dendrogasteridae  (parasitic  crustaceans  of  the 
order  Ascothoracida).]  [In  Russian.]  Uchen.  Zap.  Leningr. 
Cos.  Univ.  35(172):42-89. 
.   1957.  Dendrogasteridae  (Entomostraca,  Ascothorac- 


ida) aus  den  Asteroidea  der  Beringsee.  [In  Russian  with 
German  summary.]  Tr.    Leningr.   O-va.   Estestvoispyt. 
73(4):58-63. 
.    1970.    [Dispersal  routes  and  phylogeny  of  Ascotho- 


racida.] [In  Russian.]  Pp.  153-163  in  [Questions  on  Evolu- 
tionary Morphology  and  Biocenology].  Kazan  University 
Press,  Kazan. 
.    1976.   [Ascothoracida.]  [In  Russian.]  Kazan  Univer- 


sity Press,  Kazan.  141  p. 

YOSII,  N.    1931.    Note  on  Myriocladus.  J.  Fac.  Sci.  Imp. 
Univ.  Tokyo  Sect.  IV  Zool.  2:337-349. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  19,  pp.  455^*96,  74  figs.,  22  tables  May  14,  1982 


SMALL  TERRESTRIAL  GROUND-BEETLES  OF 

CENTRAL  AMERICA  (CARABIDAE: 

BEMBIDIINA  AND  ANILLINA) 

By 
Terry  L.  Erwin 

Department  of  Entomology,  Smithsonian  Institution, 
Washington,  D.C.  20560 


ABSTRACT:  This  paper  provides  a  synopsis  of  the  tribe  Bembidiini  with  an  overview  of  the  three  included 
subtribes,  Bembidiina,  Tachyina,  and  Anillina  for  Central  America.  Anillina  is  thought  to  be  a  polyphyletic 
grade  derived  from  genus  Paratachys  of  the  Tachyina,  but  no  formal  classificatory  action  on  this  hypothesis  is 
taken.  Based  on  structural  characters,  Horologion,  previously  classified  as  Psydrini  or  Trechini,  probably  also 
belongs  to  this  grade.  Thirteen  new  species  of  Bembidion  are  described  from  the  following  type-localities: 
purulha.  22.0  km  south  of  Purulha,  Baja  Verapaz,  Guatemala;  lavernae,  Cerro  Buenavista,  San  Jose  Province, 
Costa  Rica;  quetzal,  20.0  km  west  of  San  Pedro  Sacatepequez  at  Rio  Nahuala,  Quezaltenango,  Guatemala; 
diabola,  Volcan  Irazii,  Cartago  Province,  Costa  Rica;  edwardsi,  Cerro  Buenavista,  San  Jose  Province,  Costa 
Rica;  aeger,  Cerro  de  la  Muerte,  Cartago  Province,  Costa  Rica;  chiriqui,  8.0  km  west  of  Boquete,  Chiriqui 
Province,  Panama;  nahuala,  27.0  km  northwest  of  San  Marcos,  San  Marcos,  Guatemala;  franiae,  4.8  km  east 
of  San  Mateo  Ixtatan,  Huehuetenango,  Guatemala;  cartes,  La  Lima,  Cortes,  Honduras;  armuettes,  Puerto 
ArmueUes,  Chiriqui  Province,  Panama;  barrensis,  Barro  Colorado  Island;  ixtatan,  4.8  km  east  of  San  Mateo 
Ixtatan,  Huehuetenango,  Guatemala.  Four  new  species  of  Geocharidius  are  described  from  the  following  type- 
localities:  romeoi,  13.0  km  south  of  Purulha,  Baja  Verapaz,  Guatemala;  gimlii,  7.7  km  south  of  San  Juan 
Ixcoy,  Huehuetenango,  Guatemala;  phineus,  1.6  km  south  of  Pantin,  Baja  Verapaz,  Guatemala;  tagliantii, 
Totonicapan,  Totonicapan,  Guatemala.  The  22  Bembidion  species  presently  known  from  Central  America  are 
arrayed  in  nine  species  groups.  Corresponding  groups  for  the  five  known  Geocharidius  species  are  not  used 
because  of  the  small  size  of  the  fauna  and  their  questionable  relationships  with  poorly  known  northern  taxa. 
Distribution  and  relationships  are  discussed  in  general,  but  detailed  analysis  is  deferred  until  species-group 
taxonomy  is  undertaken  on  a  broader  geographic  scale.  Notes  are  provided  on  natural  history  and  distribution 
for  each  species;  dot  maps  illustrate  the  known  range  of  each  taxon. 

INTRODUCTION  Bates' s  papers.  The  anilline  genus  Geocharidius 

Nearly  100  years  have  passed  since  the  last  was  discussed  more  recently  by  Taglianti  (1973), 

comprehensive  paper  on  Central  American  but  his  coverage  did  not  add  new  species  for 

Bembidion  was  produced  by  H.  W.  Bates.  Dar-  Central  America.  Presently,  five  species  are 

lington  (1934)  described  a  single  species.  Other  known  from  the  area,  an  increase  of  four  since 

than  these  two  contributions,  the  genus  has  re-  Bates' s  contributions  to  Biologia   Centrali- 

ceived  no  taxonomic  attention  for  Central  Americana  (Bates  1882,  1884).  It  therefore  can 

America.  Presently,  22  species  are  known  from  be  assumed  that  after  nearly  100  years  the  Bem- 

the  area,  an  increase  of  17  since  the  time  of  bidion  fauna  of  Central  America  (defined  here 

[455] 


456 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


as  the  mainland  and  coastal  islands  from  Gua- 
temala to  Panama)  is  still  not  completely  known; 
the  anilline  fauna  is  essentially  untouched,  prob- 
ably a  result  of  their  living  in  deep  soil  humus, 
thus  hard  to  collect  except  by  special  means. 
The  tachyine  fauna,  here  discussed  at  the  ge- 
neric level  only,  is  huge  (Erwin  1974b)  and  is 
presently  under  revision  (e.g.,  Erwin  1973, 
1974a,  1975,  1977). 

In  order  to  provide  data  for  a  series  of  studies 
now  underway  by  D.  H.  Kavanaugh  and  me  on 
the  genus  Bembidion  (Erwin  and  Kavanaugh 
1980,  1981),  I  am  publishing  here  an  extract  from 
my  "Ground  Beetles  of  Central  America"  in 
advance  of  the  volume  in  which  it  belongs.  The 
present  study  is  a  faunal  documentation  which 
will  provide  data  for  future  analyses.  Comments 
pertaining  to  biogeography  are  given  below,  but 
since  the  Central  American  bembidiine  fauna 
represents  the  "tail-ends"  of  more  northernly  or 
southernly  distributed  groups,  phylogenetic  and 
detailed  biogeographic  analyses  will  be  dis- 
cussed elsewhere  (e.g.,  Erwin  and  Kavanaugh 
1981). 

MATERIALS 

This  paper  is  based  on  examinations  of  1002 
specimens  of  Bembidion  and  44  specimens  of 
Geocharidius  from  Central  America,  as  well  as 
numerous  specimens  of  other  Bembidiini  from 
other  parts  of  the  world.  Following  is  a  list  of 
depositories  for  these  materials;  acronyms 
based  on  Erwin  (1970)  are  given  preceding  each 
entry:  CAS — California  Academy  of  Sciences, 
San  Francisco,  California  94118,  D.  H.  Kava- 
naugh; MCZ — Museum  of  Comparative  Zoolo- 
gy, Cambridge,  Massachusetts,  M.  Thayer,  A. 
Newton;  MNHP — Museu  National  d'Histoire 
Naturelle,  Paris,  H.  Perrin,  J.  Menier;  UASM— 
Strickland  Museum,  University  of  Alberta,  Ed- 
monton, Canada,  G.  E.  Ball;  BMNH— British 
Museum  (Natural  History),  London,  P.  Ham- 
mond, M.  Bacchus;  HPST— Canal  Zone,  Henry 
P.  Stockwell;  AMNH — American  Museum  of 
Natural  History,  New  York,  L.  Herman; 
FMNH — Field  Museum  of  Natural  History, 
Chicago,  H.  Dybas;  FDAG— Florida  Depart- 
ment of  Agriculture  Insect  Collection,  Gaines- 
ville, Florida,  R.  Woodruff;  UMAA — Universi- 
ty of  Michigan,  Ann  Arbor,  R.  D.  Ward; 
UAIC — University  of  Arkansas,  Fayetteville, 
R.  T.  Allen;  GRNO— Milwaukee  Public  Mu- 


seum, Wisconsin,  G.  R.   Noonan;  JNEG — 78 
Versailles,  9,  Bb  De  Lesseps,  France,  J.  Negre. 

METHODS 

General  procedural  methods  are  as  I  used  be- 
fore (Erwin  1970,  1973,  1974a,  1975;  Erwin  and 
Kavanaugh  1981).  Measures  for  various  body 
parts  are  coded  as  follows  and  are  presented  in 
the  species  descriptions  as  single  specimen  mea- 
sures, if  that  is  all  I  saw,  or  ranges  based  on  the 
smallest  and  largest  of  all  specimens  studied — 
all  specimens  were  measured  with  an  electronic 
measuring  device  (Erwin  1978)  and  measures  are 
presented  in  millimeters.  ABL  =  apparent  body 
length,  that  length  used  by  most  previous  au- 
thors as  total  length  as  measured  by  holding  up 
a  ruler  alongside  the  specimen  (see  Erwin  and 
Kavanaugh  1981).  SBL  =  standardized  body 
length,  that  length  introduced  by  Ball  (1972)  and 
modified  by  Kavanaugh  (1979),  and  equal  here 
to  the  sum  of  LH,  LP,  and  LE.  TW  =  total 
width  across  the  widest  portion  of  the  elytra, 
actually  measured  as  the  left  elytron  (WE)  and 
doubled  to  obtain  value.  LH  =  length  of  head 
from  front  edge  of  clypeus  to  base  of  head  at 
point  of  cervical  constriction  and  taken  along 
midline.  LP  =  length  of  pronotum  from  base  to 
apex  along  midline.  LE  =  length  of  elytron, 
longest  one,  along  suture  from  apex  of  scutellum 
to  apex  of  sutural  projection.  WE  =  width  of 
elytron  at  widest  portion.  All  label,  specimen 
measurement,  and  field  data  about  specimens 
are  computerized  (L.  J.  M.  Erwin  1976),  and 
these  records  formed  the  base  for  study  and 
analysis.  Data  concerning  material  examined  are 
presented  in  tables  and  maps;  tables  summarize 
data  deduced  from  specimen  labels,  however, 
localities  are  presented  as  latitude  and  longitude 
except  in  the  case  of  types  where  place  names 
are  also  used.  Elevations  are  given  in  meters 
even  though  specimen  labels  may  have  them  in 
English  units;  occasionally,  specimen  labels  give 
imprecise  altitudinal  data  or  none  at  all,  thus,  in 
these  cases  tables  include  the  abbreviation  'ca.' 
to  indicate  approximate  altitude  derived  from 
topographic  maps.  Chaetotaxy  reference  follows 
that  of  Erwin  (1974a)  and  the  term  'interneur,' 
used  for  the  structure  between  elytral  intervals, 
is  described  in  Erwin  (1974a).  Microsculpture 
micrographs  were  made  only  for  species  where 
sufficient  study  material  allowed  coating  of  body 
parts. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


457 


FIGURE  1.     Habitus  of  Bembidion  vernale  Bates,  dorsal  aspect,  female  from  Cerro  Punta,  Chiriqui,  Panama. 


458 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


TRIBE  BEMBIDIINI 

The  bembidiines  are  a  diverse  lot  of  mostly 
small  beetles  distributed  worldwide  and  whose 
primary  apotypic  feature  is  the  subulate  apical 
palpomere.  The  tribe  is  represented  by  three 
subtribes  in  Central  America:  Bembidiina, 
Tachyina,  and  Anillina. 

Bembidiinas  are  much  larger,  as  a  rule,  than 
members  of  the  other  subtribes,  often  are  col- 
orful, almost  always  have  large  eyes,  and  do  not 
possess  an  elytral  recurrent  groove  (however, 
the  apex  of  interneur  5  is  often  depressed).  Bem- 
bidion  includes  hundreds,  if  not  thousands,  of 
species  worldwide,  but  most  of  these  are  dis- 
tributed in  the  Holarctic  and  South  Temperate 
regions.  In  Central  America,  there  are  only  22 
species  known,  but  undoubtedly  some  more-al- 
ticolous  species  will  be  discovered  with  addi- 
tional collecting  in  remote  upland  areas.  These 
beetles  live  in  such  diverse  habitats  as  sea 
beaches  and  mountain  tops  above  timber  line. 
They  are  particularly  common  and  diverse  along 
gravelly  streams  and  rivers  and  in  upland  forest 
above  2000  meters.  The  major  taxonomic  prob- 
lem remaining  in  this  genus  is  to  segregate  and 
define  its  components  on  a  worldwide  basis.  Ne- 
tolitzky  (1942,  1943)  and  Lindroth  (1963)  provid- 
ed an  excellent  foundation  with  which  to  start; 
D.  H.  Kavanaugh  and  I  have  taken  up  the  task. 
The  former  authors  have  shown  beyond  doubt 
that  male  genitalic  characteristics  are  of  great 
value  in  several  Holarctic  groups  for  determin- 
ing relationships.  Erwin  and  Kavanaugh  (1981) 
showed  the  same  to  be  true  for  the  female  re- 
productive system.  Reclassification  of  this  com- 
plex and  diverse  genus  will  require  a  world  over- 
view, reevaluation  of  character  systems,  and 
close  scrutiny  of  geographic  ranges.  Much  more 
collecting  will  need  to  be  done,  especially  in  the 
western  United  States,  South  American  Andes, 
and  Palearctic  Asia.  The  larval  stages  of  many 
groups  of  Bembidiina  are  known  (see  Lindroth 
1963);  these  will  aid  systematic  studies  because 
taxonomically  useful  characters  are  present  (Er- 
win 1975). 

Tachyinas  are  small  to  medium-sized  bembi- 
diines with  much  greater  diversity  and  richness 
than  bembidiinas.  Tachyinas  almost  always 
have  eyes,  many  are  colorful  (some  metallic), 
and  most  possess  a  recurrent  groove  near  the 
elytral  apex.  Most  Central  American  genera  are 
composed  of  numerous  species.  This  subtribe 


constitutes  one  of  the  two  largest  groups  of  the 
Central  American  fauna  (the  other  being  Ago- 
nina,  Platynus,  and  its  relatives).  Tachyina  bee- 
tles live  in  diverse  habitats,  from  intertidal  zones 
to  rain  forest  canopy;  they  are  extremely  diverse 
along  riparian  systems,  and  some  have  adapted 
to  living  with  ants.  They  are  a  lowland  faunal 
component  and  have  not  become  true  alticoles 
as  have  Bembidion  and  other  carabid  groups 
such  as  the  trechines,  indicating  that  they  are  a 
later  taxon  pulse  element  of  the  fauna  than  are 
the  Bembidiina  (Erwin  1979,  1981;  Erwin  and 
Adis  1982).  Erwin  (1973,  1974a,  1974b,  1975)  has 
begun  a  world  revisionary  study  of  this  group  of 
beetles.  As  with  bembidiinas,  reclassification  of 
this  complex  and  diverse  tribe  will  require  a 
world  overview,  reevaluation  of  character  sys- 
tems, and  close  scrutiny  of  geographic  ranges. 
Much  more  collecting  will  need  to  be  done,  es- 
pecially in  the  tropics.  The  larval  stages  of 
Tachyina  are  mostly  unknown  (see  Erwin  1975). 
Anillinas  are  minute,  apterous,  blind  beetles 
which  inhabit  deep  humus  in  upland  forests  and 
are  also  found  in  caves  and  fissures.  Difficulty 
in  finding  these  beetles  prevents  accumulation 
of  enough  material  to  adequately  revise  most 
genera.  Undoubtedly,  more  species  will  be  dis- 
covered in  upland  forests  by  use  of  special  col- 
lecting techniques  (e.g.,  Berlese  funnel).  Anilli- 
nas, as  currently  classified,  are  easily  recognized 
by  their  small  size,  pale  coloration,  and  lack  of 
eyes.  Some  cave-dwelling  Ardistomina  (Scari- 
tini)  have  also  lost  eyes,  but  remain  easily  rec- 
ognized by  body  shape  (pedunculate).  Some  tre- 
chines in  Middle  America  have  also  adapted  to 
caves  and  become  blind,  but  they  can  be  im- 
mediately recognized  by  their  long  apical  pal- 
pomeres  and  sulcate  head.  There  are  a  few  wing- 
less, pale,  nearly  blind  Mioptachys  (Tachyina) 
in  the  same  habitat  with  Geocharidius  (the  only 
genus  of  Central  American  Anillina);  the  former 
always  have  some  eye  facets,  however,  and  pos- 
sess a  deeply  engraved  and  sulcate  recurrent 
groove  on  the  lateroapical  portion  of  the  elytron. 
Jeannel  (1937,  1963a,  1963b)  attempted  a  phy- 
logenetic  and  biogeographic  synthesis  of  this 
group,  however,  he  was  partially  thwarted  by 
lack  of  material,  his  assumption  that  the  group 
is  monophyletic,  and  the  'reduction'  of  most 
character  suites  leaving  poorly  defined  taxa.  Re- 
cent discovery  of  an  apparently  eyed  Anillina 
from  lowland  Brazil  suggests  a  relationship  of 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


459 


the  subtribe  to  the  tachyina  group  "Polyderi- 
dius."  Possibly,  Anillina  represents  a  grade  of 
numerous  parallel  lineages  derived  from  Para- 
tachys  and  allies,  each  of  which  adapted  inde- 
pendently to  deep-humus  environments.  This 
possibility  would  explain  several  puzzles.  For 
example,  all  anillinas  have  the  anterior  tibia 
notched  obliquely — an  apotypic  trend  found 
among  all  the  higher  tachyinas,  which  reaches 
its  ultimate  development  in  Micratopus  and 
Lymnastis.  Also  explained  would  be  the  occur- 
rence of  these  small,  blind,  wingless  beetles  on 
remote  islands  and  all  continents  (except  Ant- 
arctica). At  present,  the  only  explanation  is  that 
they  are  an  ancient  group  predating  the  initial 
rift  of  Pangaea.  This  explanation  is  not  concor- 
dant with  many  other  aspects  of  carabid  evolu- 
tion and  radiation.  It  seems  to  me  that  a  poly- 
phyletic  origin  hypothesis  now  best  fits 
observations,  and  Paratachys  and  allies  are  the 
source  of  the  anillina  grade,  as  they  are  also  the 
probable  source  of  "Lymnastine"  radiation. 

Horologion,  a  monotypic  genus  (known  from 
only  one  specimen)  from  West  Virginia  repre- 
sented by  H.  speokoites,  has  long  troubled  clas- 
sifiers (Valentine  1932;  van  Emden  1936;  Ball 
1963).  Considering  the  evidence  presented 
above  and  the  fact  that  many  character  states 
are  shared  among  Horologion  and  Anillinas,  and 
especially,  that  these  share  the  derived  tibial 
configuration  mentioned  above,  I  here  suggest 
that  Horologion  belongs  in  the  anillina  grade 
and  that  its  origin  lies  in  the  Tachyina. 

For  recent  treatments  of  Bembidiini  or  its 
components  see  Basilewsky  (1968,  1972 — Mad- 
agascan  and  St.  Helena  Bembidiini),  Darlington 
(1962— New  Guinea  Bembidiini),  Erwin  (1973, 
1974a,  1974b,  1975,  1976,  1977— Tachyina),  Er- 
win and  Kavanaugh  (1980,  1981— North  Amer- 
ican Bembidion),  Kryzhanovskiy  (1970 — 
U.S.S.R.  Tachyina),  Lindroth  (1963,  1966,  1975, 
1976,  1980 — North  American,  Australian,  and 
Asian  Bembidiini),  de  Mire  (1976 — African 
Tachyina),  Pawlowski  (1974— Polish  Bembidi- 
ini). 

Genus  Bembidion  Latreille 

Bembidium  LATREILLE,  1802:82.  [Type-species  Cicindela 
quadrimaculata  Linne,  1761:211,  subsequent  designation 
by  Andrewes(1935:17).] 

For  detailed  generic  synonymies  see  An- 
drewes  (1935:92),  Ball  (1963),  Lindroth 


(1963:206),  Muller  (1918),  and  Netolitzky  (1942, 
1943). 

DIAGNOSTIC  COMBINATION. — Small  to  medi- 
um-sized, more  or  less  depressed  beetles  with 
broad  elytra,  narrow  forebody,  and  long  slender 
appendages.  Color  various,  often  with  metallic 
luster,  elytra  sometimes  spotted.  Terminal  pal- 
pomere  subulate;  eyes  moderate  to  large,  anten- 
nomeres  3-11  entirely  pubescent,  mentum  with 
well-developed  tooth;  mandibles  unisetose. 
Pronotum  with  two  midlateral  and  usually  two 
basilateral  setae;  elytron  with  two  or  more  discal 
setae  in  third  interval  or  adjacent  interneur. 
Elytron  without  recurrent  groove,  although  apex 
of  fifth  interneur  usually  sulcate.  Microsculpture 
various.  Female  stylus  small,  spatulate,  unise- 
tose. Male  parameres  2-  or  3-setose.  Flight 
wings  present,  absent,  or  dimorphic;  reduction 
in  size  and  length  of  various  degrees.  ABL  = 
2.43-6.11  mm  in  Central  American  species. 

NATURAL  HISTORY. — Species  of  this  genus 
live  on  sea  beaches,  along  streams  and  rivers, 
and  in  forests;  they  can  be  found  during  their 
quiescent  periods  under  leaf  litter,  logs,  or 
stones.  Several  species  in  Central  America  are 
true  alticoles;  most  of  these  are  wingless.  The 
larvae  of  many  European  and  some  North 
American  species  are  known  and  described,  but 
none  in  Central  America  have  been  discovered 
in  either  larval  or  pupal  stages.  Adults  of  Bem- 
bidion species  are  carnivorous;  in  addition  to 
small  arthropods,  they  also  eat  arthropod  eggs 
(Frank  1971).  Their  usual  life  cycle  requires  one 
year;  larvae  are  present  in  the  fauna  in  summer 
for  the  most  part,  however,  some  species  over- 
winter in  this  stage  in  the  north  (Lindroth  1963). 
Bembidion  species  are  dominant  members  of 
most  northern  riparian,  alpine,  and  lake  shore 
communities.  In  the  tropics,  this  dominance 
does  not  exist  at  lower  elevations,  however,  I 
have  seen  it  in  upland  riparian  habitats  in  Pan- 
ama and  in  the  Subparamo  habitat  in  Costa  Rica. 

NOTES. — Bembidion  of  the  southern  United 
States,  Mexico,  and  South  America  are  in  need 
of  a  detailed  taxonomic  study.  Therefore,  I  have 
restricted  coverage  here  to  only  Central  Amer- 
ica within  the  limits  set  forth  in  the  Introduction. 
The  North  American  Bembidion  fauna  consists 
of  more  than  200  species  (Lindroth  1967),  that 
of  Central  America  22!  This  drastic  reduction  in 
species  numbers  occurs  in  Mexico,  most  dra- 
matically at  the  southern  edge  of  the  Transverse 


460 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


FIGURE  2.  Elytron,  left,  dorsal  aspect  of  (a)  Bembidion 
rogersi  Bates,  alate  form,  7.5  km  south  of  Ojo  de  Agua,  Car- 
tago,  Costa  Rica,  male;  (b)  same,  brachypterous  form,  same 
locale,  male. 


Volcanic  Belt  and  Isthmus  of  Tehuantepec. 
From  this  line  southward,  Bembidion  is  never 
as  diverse  as  it  is  in  North  America,  although 
species  are  found  as  far  south  as  Tierra  del  Fue- 
go  (Darlington  1965). 

The  Bembidion  fauna  of  North  America  is 
very  complex  and  difficult  to  work  with  because 
of  many  closely  related  species  in  most  species 
groups.  That  of  Central  America,  however,  is 
quite  the  contrary  because  there  are  only  a  few 
species  in  each  of  the  nine  represented  groups, 
the  tail-ends  of  otherwise  diverse  northern  and 
southern  groups.  Male  genitalic  character  states 
are  excellent  for  delimiting  species  and  species 
groups,  female  genitalic  characteristics  for 
species  groups;  both  these  suites  usually  corre- 
late well  with  external  structure  (Erwin  and  Ka- 
vanaugh  1981)  which  then  can  be  utilized  for 
diagnostic  features.  Externally,  the  beetles  dif- 
fer in  states  of  the  frontal  furrows,  metasternal 
process,  dorsal  elytral  chaetotaxy  and  structure 
of  the  interneurs,  microsculpture,  and  pronotal 
shape,  even  though  the  beetles  themselves  are 
similar  in  habitus.  In  Central  America,  two  hab- 
itus types  occur:  the  winged  type  (Fig.  2a)  with 
large  square  humeri  and  narrow  forebody  and 
head,  and  the  brachypterous  type  (Fig.  2b)  with 
rounded  and  narrow  humeri,  arcuate-sided  ely- 
tra, and  more  robust  forebody  and  head.  These 
two  types  occur  among  different  individuals  of 
the  same  population  of  wing-dimorphic  species 
such  as  B.  rogersi  and  B.  nahuala. 

Adults  can  easily  be  sexed  in  two  ways.  Males 
have  two  basal  tarsomeres  of  the  anterior  leg 
dilated  and  with  modified  setae  beneath,  forming 
a  loosely  biserrate  "spongy"  pad,  and  two  setae 
along  the  caudal  edge  of  sternum  VI;  females 
have  no  "spongy"  pad  or  dilated  tarsomeres  and 
have  four  setae  on  sternum  VI. 


ARRANGEMENT  OF  TAXA:  BEMBIDION. — I  here 
follow  Lindroth's  arrangement  of  species  groups 
and  add  the  rogersi,  vernale,  nahuala,  franiae, 
and  cognatum  groups,  none  of  which  were  cov- 
ered by  him.  Arrangement  of  taxa  within  groups 
is  based  on  complexity  of  male  genital  structure, 
from  simple  to  complex.  Athough  this  may  not 
be  phylogenetically  accurate,  it  suits  the  pur- 
pose of  this  paper  in  its  attempt  to  simply  doc- 
ument the  fauna  for  further  study.  Such  further 
study  must  necessarily  be  based  on  revisionary 
work  with  a  much  broader  geographic  base  than 
Central  America. 

CHECKLIST  OF  BEMBIDION  SPECIES  OF  CEN- 
TRAL AMERICA. 

The  semistriatum  group 

1.  B.  purulha,  new  species 
The  rogersi  group 

2.  B.  rogersi  Bates,  1878:602 
The  vernale  group 

3.  B.  mexicanum  Dejean,  1831:126 

4.  B.  vernale  Bates,  1882:149 

5.  B.  satellites  Bates,  1884:291 

6.  B.  lavernae,  new  species 

7.  B.  quetzal,  new  species 

8.  B.  diabola,  new  species 

9.  B.  vulcanium  Darlington,  1934:157 

10.  B.  edwardsi,  new  species 

11.  B.  aeger,  new  species 

12.  B.  chiriqui,  new  species 
The  nahuala  group 

13.  B.  nahuala,  new  species 
The  franiae  group 

14.  B.  franiae,  new  species 
The  incrematum  group 

15.  B.  aratum  LeConte,  1852:189 
The  dorsale  group 

16.  B.  cartes,  new  species 

17.  B.  sparsum  Bates,  1882:151 

18.  B.  armuelles,  new  species 
The  affine  group 

19.  B.  barrensis,  new  species 
The  championi  group 

20.  B.  cyclodes  Bates,  1884:290 

21.  B.  championi  Bates,  1882:148 
The  cognatum  group 

22.  B.  ixtatan,  new  species 

Key  to  Species  of  Bembidion  of 
Central  America 

1 .  Pronotum  with  hind  angles  complete- 
ly rounded,  base  markedly  lobed  as  in 
Figure  21 2 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


461 


TABLE  1.     NUMBERS  AND  PERCENTAGES  OF  WINGED  VERSUS  NON-WINGED  INDIVIDUALS  IN  THREE  POPULATIONS  OF  (A) 
Bembidion  rogersi  BATES  AND  (B)  Bembidion  nahuala,  NEW  SPECIES. 


-  Wing 
female 

+  Wing 
female 

%  Female 
-  wing 

-  Wing 
male 

+  Wing 
male 

%  Male 
-  wing 

%  Total 
-  wing 

Locality 

(A) 

Cerro  de  la  Muerte 

54 

7 

88% 

99 

20 

83% 

85% 

Volcan  Irazii 

4 

4 

50% 

7 

1 

87% 

69% 

Volcan  de  Chiriqui 

6 

4 

60% 

3 

2 

60% 

60% 

(B) 

Rio  Nahuala 

5 

1 

83% 

0 

0 

— 

83% 

San  Marcos 

26 

9 

74% 

16 

2 

89% 

79% 

Cerro  Zunil 

1 

0 

100% 

0 

0 

— 

100% 

Finca  San  Rafael 

1 

0 

100% 

0 

0 

— 

100% 

Pronotum  with  hind  angles  acute  or 
square,  base  not  lobed  3 

2(1).  Elytron  with  interneur  7  absent,  in- 
terneur  6  extended  to  at  least  middle 

19.  B.  cyclodes  Bates 

Elytron  with  interneurs  6  and  7  pres- 
ent, short,  not  extended  beyond  basal 
sixth 20.  B.  champion!  Bates 

3(1).  Frontal  furrows  doubled  on  clypeus 
and  behind  posterior  supraorbital 
seta,  furrows  moderately  or  markedly 
convergent;  humerus  somewhat  paler 
than  disc  of  elytron;  small  beetles, 

less  than  3.0  mm  4 

Frontal  furrows  not  doubled,  not  con- 
vergent; humerus  various;  larger  bee- 
tles, usually  more  than  3.0  mm  5 

4(3).  Frontal  furrows  deeply  impressed  and 
convergent,  doubled  throughout  their 

length  22.  B.  ixtatan,  new  species 

Frontal  furrows  shallow  and  single 
between  eyes,  nearly  parallel  except 
on  clypeus  where  they  converge  and 

are  doubled 

19.  B.  barrensis,  new  species 

5(3).       Metasternal  process  between  meso- 

coxae  entirely  unmargined 6 

Metasternal  process  clearly  margined, 
margin  removed  from  apex  and 
abruptly  or  gradually  raised 8 

6(5).       Dorsal  surface  without  microsculp- 

ture,  surfaces  shiny   

13.  B.  nahuala,  new  species 

Dorsal  surface  with  well-developed 
microsculpture,  surfaces  dull  7 


7(6).  Elytron  bifoveate,  foveae  around  se- 
tae deep  and  wide,  each  extended 
across  about  three  intervals;  elytral 
microsculpture  of  wide  meshes,  on 
head  and  pronotum  variously  effaced 
or  nearly  isodiametric 

2.  B.  rogersi  Bates 

Elytron  bipunctate,  punctures  around 
dorsal  setae  deep  and  small,  not  ex- 
tended beyond  middle  of  adjacent  in- 
terval; elytral  microsculpture  of  finely 
impressed  lines  or  wide  meshes,  sur- 
face subiridescent,  on  head  clearly 
and  evenly  isodiametric,  slightly  gran- 
ular, more  evenly  transverse  and  less 
impressed  on  pronotum 

14.  B.  franiae,  new  species 

8(5).  Elytron  with  setae  Ed3  and  Ed5  in  in- 
terval 3  and  not  touching  adjacent  in- 
terneurs    9 

Elytron  with  these  setae  in  or  touch- 
ing interneur  3 12 

9(8).       Elytron  without  microsculpture, 

markedly  shiny  

15.  B.  aratum  LeConte 

Elytron  with  perfectly  isodiametric 
and  well-engraved  microsculpture, 
surfaces  dull  10 

10(9).  Pronotum  (Fig.  19)  narrow,  about  as 
wide  as  head  across  eyes;  length  3.47 
to  4.23  mm  ....  17.  B.  sparsum  Bates 
Pronotum  broad,  much  wider  than 
head  across  eyes;  length  3.75  to  4.41 
mm  1 1 

11(10).  Pronotum  (Fig.  20)  markedly  convex 


462  PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 

TABLE  2.     VARIATION  IN  TOTAL  LENGTH  (SBL  in  mm)  WITHIN  AND  AMONG  Bembidion  SPECIES  OF  CENTRAL  AMERICA. 


Sex 

(») 

Minimum 

Maximum 

Mean 

±SE 

SD 

CV 

01.  B. 

purulha 

m 
f 

1 

1 

4.19 

4.53 

4.19 

4.53 

— 

— 

— 

— 

02.  B. 

rogersi 

m 
f 

135 
88 

3.78 
3.60 

4.26 
4.63 

4.25 
4.20 

0.02 
0.02 

0.22 
0.24 

5.24 
5.69 

03.  B. 

mexicanum 

m 
f 

33 
29 

5.04 
4.95 

6.11 
5.95 

5.50 
5.41 

0.05 
0.05 

0.29 
0.26 

5.36 
4.88 

04.  B. 

vernale 

m 

f 

182 
113 

4.39 
4.43 

5.64 
5.66 

5.06 
5.14 

0.02 
0.02 

0.24 
0.24 

4.72 
4.68 

05.  B. 

satellites 

m 
f 

30 

25 

4.20 

4.22 

5.12 
5.17 

4.61 
4.79 

0.04 
0.05 

0.24 
0.23 

5.25 
4.87 

06.  B. 

lavernae 

m 
f 

X 
1 

4.18 

4.18 







— 

07.  B. 

quetzal 

m 

f 

9 
18 

4.74 
4.77 

5.59 

5.53 

5.21 
5.23 

0.09 
0.06 

0.27 
0.24 

5.26 
4.61 

08.  B. 

diabola 

m 

f 

31 

22 

3.56 
3.64 

4.38 
4.49 

3.71 
4.10 

0.04 
0.05 

0.22 
0.24 

5.47 
5.93 

09.  B. 

vulcanium 

m 
f 

18 
15 

3.73 
3.11 

4.49 
4.36 

4.12 
4.02 

0.05 
0.08 

0.20 
0.32 

4.75 
7.85 

10.  B. 

edwardsi 

m 

f 

X 

3 

3.77 

4.00 

3.87 

0.07 

0.12 

3.09 

11.  B. 

aeger 

m 

f 

18 
16 

3.05 
3.26 

3.74 
3.75 

3.49 
3.44 

0.04 
0.03 

0.18 
0.12 

4.19 
3.38 

12.  B. 

chiriqui 

m 
f 

34 
10 

3.42 
3.68 

4.15 
4.25 

3.84 
3.98 

0.03 
0.05 

0.19 
0.17 

4.89 

4.22 

13.  B. 

nahuala 

m 
f 

19 
41 

2.87 
3.04 

3.77 
4.00 

3.24 
3.50 

0.06 
0.03 

0.27 
0.21 

8.28 
5.93 

14.  B. 

franiae 

m 
f 

1 
1 

4.12 
3.98 





— 



— 

15.  B. 

aratum 

m 
f 

2 
3 

4.03 
4.24 

5.03 
4.48 

4.53 
4.35 

0.50 
0.07 

0.71 
0.12 

15.61 

2.77 

16.  B. 

cones 

m 
f 

10 

22 

3.75 
3.75 

4.41 
4.40 

4.17 
4.13 

0.04 
0.04 

0.13 
0.19 

3.22 
4.67 

17.  B. 

sparsum 

m 
f 

9 
8 

3.47 
3.52 

4.10 
4.23 

3.74 
3.82 

0.08 
0.08 

0.23 
0.24 

6.16 
6.20 

18.  B. 

armuelles 

m 
f 

X 

1 

4.32 

4.32 

— 

— 





19.  B. 

barrensis 

m 
f 

1 
X 

2.94 

2.94 

I 

— 

— 



20.  B. 

cyclodes 

m 

f 

X 

1 

3.05 

3.05 





— 

— 

21.  B. 

championi 

m 
f 

X 
X 

— 





— 

— 



22.  B. 

ixtatan 

m 
f 

3 
4 

2.43 
2.54 

2.90 

2.85 

2.70 
2.69 

0.01 
0.07 

0.24 
0.14 

9.03 
5.06 

"X"  =  sex  not  available  for  measurement. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


463 


TABLE  3.     VARIATION  IN  TOTAL  WIDTH  (TW  in  mm)  WITHIN  AND  AMONG  Bembidion  SPECIES  OF  CENTRAL  AMERICA. 


Sex 

(«) 

Minimum 

Maximum 

Mean 

±SE 

SD 

CV 

01.  B 

purulha 

m 

1 

1.79 

1.79 

— 

— 

— 



f 

1 

1.82 

1.82 

— 

— 

— 

— 

02.  B 

rogersi 

m 

135 

1.58 

2.13 

1.82 

0.01 

0.22 

5.24 

f 

88 

1.57 

1.98 

1.80 

0.01 

0.11 

5.93 

03.  B 

mexicanum 

m 

33 

2.07 

2.80 

2.44 

0.02 

0.14 

5.62 

f 

29 

2.09 

2.80 

2.42 

0.02 

0.13 

5.52 

04.  B 

vernale 

m 

182 

1.89 

2.61 

2.18 

0.01 

0.12 

5.45 

f 

113 

1.80 

2.51 

2.22 

0.01 

0.12 

5.47 

05.  B 

satellites 

m 

30 

1.64 

2.12 

1.96 

0.02 

0.11 

5.50 

f 

26 

1.77 

2.30 

2.04 

0.02 

0.11 

5.18 

06.  B 

lavernae 

m 

X 

— 

— 

— 

— 

— 

— 

f 

1 

1.78 

1.78 

— 

— 

— 

— 

07.  B 

quetzal 

m 

9 

2.04 

2.38 

2.24 

0.04 

0.11 

4.72 

f 

18 

2.08 

2.45 

2.24 

0.03 

0.13 

5.72 

08.  B 

diabola 

m 

31 

1.52 

1.89 

1.74 

0.02 

0.10 

5.59 

f 

22 

1.52 

1.93 

1.76 

0.02 

0.11 

6.26 

09.  B 

vulcanium 

m 

18 

1.63 

2.00 

1.89 

0.02 

0.08 

4.48 

f 

15 

1.58 

2.04 

1.89 

0.03 

0.13 

6.67 

10.  B 

edwardsi 

m 

X 













f 

3 

1.59 

1.66 

1.62 

0.02 

0.04 

2.34 

11.  -B 

aeger 

m 

18 

1.28 

.65 

1.51 

0.02 

0.09 

5.82 

f 

16 

1.34 

.81 

1.50 

0.03 

0.11 

7.45 

12.  £ 

chiriqui 

m 

34 

1.54 

.83 

1.70 

0.01 

0.08 

4.58 

f 

10 

1.65 

.94 

1.77 

0.03 

0.09 

5.25 

13.  B 

nahuala 

m 

19 

1.20 

.78 

1.45 

0.03 

0.16 

11.18 

f 

41 

1.28 

.94 

1.58 

0.02 

0.12 

7.40 

14.  B 

franiae 

m 

1 

1.85 

1.85 

— 

— 

— 

— 

f 

1 

1.83 

1.83 

— 

— 

— 

— 

15.  B 

aratum 

m 

2 

1.68 

1.98 

1.83 

0.15 

0.21 

11.59 

f 

3 

1.62 

1.85 

1.77 

0.08 

0.13 

7.21 

16.  B 

cartes 

m 

10 

3.98 

4.14 

4.17 

0.02 

0.07 

3.79 

f 

22 

1.65 

1.99 

1.84 

0.02 

0.08 

4.29 

17.  B 

sparsum 

m 

9 

1.50 

1.74 

1.62 

0.03 

0.09 

5.42 

f 

8 

1.53 

1.89 

1.71 

0.05 

0.13 

7.84 

18.  B 

armuelles 

m 

X 

— 

— 

— 

— 

— 

— 

f 

1 

1.80 

1.80 

— 

— 

— 

— 

19.  B 

barrensis 

m 

1 

1.21 

1.21 

— 

— 

— 

— 

f 

X 

— 

— 

— 

— 

— 

— 

20.  B 

cyclodes 

m 

X 

— 

— 

— 

— 

— 

— 

f 

1 

1.39 

1.39 

— 

— 

— 

— 

21.  B 

championi 

m 

X 

— 

— 

— 

— 

— 

— 

f 

X 

— 

— 

— 

— 

— 

— 

22.  B 

ixtatan 

m 

3 

1.20 

1.26 

1.24 

0.02 

0.03 

2.60 

f 

4 

1.07 

1.21 

1.14 

0.03 

0.06 

5.21 

"X"  =  sex  not  available  for  measurement. 


464 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


TABLE  4.     VARIATION  IN  TOTAL  DEPTH  (TD  in  mm)  WITHIN  AND  AMONG  Bembidion  SPECIES  OF  CENTRAL  AMERICA. 


Sex 

(n) 

Minimum 

Maximum 

Mean 

±SE 

SD 

CV 

01.  B 

purulha 

m 
f 

1 
1 

1.04 
1.12 

1.04 
1.12 

I 

~ 

— 

— 

02.  B 

rogersi 

m 
f 

135 
88 

0.81 
0.74 

1.19 
1.16 

0.95 
0.% 

0.01 
0.01 

0.08 
0.09 

8.49 
9.02 

03.  B 

mexicanum 

m 

f 

33 
29 

1.18 
1.15 

1.54 
1.47 

1.33 
1.31 

0.02 
0.01 

0.10 
0.08 

7.17 
6.14 

04.  B 

vernale 

m 

f 

182 
113 

0.86 
0.89 

1.40 
1.37 

1.10 
1.10 

0.01 
0.01 

0.01 
0.10 

8.63 
9.32 

05.  B. 

satellites 

m 
f 

30 
26 

0.87 
0.88 

1.21 
1.21 

1.04 
1.03 

0.01 
0.02 

0.07 
0.09 

7.04 
8.26 

06.  B. 

lavernae 

m 
f 

X 

1 

0.83 

0.83 

— 

— 

— 

— 

07.  B. 

quetzal 

m 
f 

9 
18 

1.01 
0.94 

.38 
.39 

1.19 
1.16 

0.04 
0.03 

0.12 
0.11 

10.37 
9.50 

08.  B. 

diabola 

m 
f 

31 

22 

0.76 
0.82 

.15 
.20 

0.96 
0.% 

0.02 
0.02 

0.10 
0.10 

9.88 

10.17 

09.  B. 

vulcanium 

m 
f 

18 
15 

0.86 

0.78 

.14 
.09 

1.00 
0.% 

0.02 
0.02 

0.08 
0.09 

7.51 
8.90 

10.  B. 

edwardsi 

m 

f 

X 

3 

0.84 

0.87 

0.85 

0.01 

0.02 

1.79 

11.  B. 

aeger 

m 
f 

18 
16 

0.52 
0.65 

0.84 
0.88 

0.73 
0.76 

0.02 
0.02 

0.09 
0.07 

12.11 
9.18 

12.  B. 

chiriqui 

m 
f 

34 
10 

0.80 

0.78 

1.05 
1.03 

0.90 
0.90 

0.01 
0.03 

0.06 
0.09 

7.00 
10.15 

13.  B. 

nahuala 

m 
f 

19 

41 

0.60 
0.64 

0.95 
0.95 

0.72 
0.81 

0.02 
0.01 

0.10 
0.07 

13.97 
8.52 

14.  B. 

franiae 

m 
f 

1 

1 

0.93 
0.85 

0.93 
0.85 

— 

— 

— 



15.  5. 

aratum 

m 
f 

2 
3 

0.92 
0.82 

0.94 
0.89 

0.93 
0.86 

0.01 
0.02 

0.01 
0.04 

1.52 
4.39 

16.  B. 

cones 

m 

f 

10 

22 

0.92 
0.84 

1.12 
1.13 

1.00 
1.00 

0.02 
0.02 

0.07 
0.07 

6.81 
7.36 

17.  B. 

sparsum 

m 
f 

9 

8 

0.76 
0.89 

0.98 

1.02 

0.89 
0.95 

0.02 
0.02 

0.07 
0.05 

8.06 
4.88 

18.  B. 

armuelles 

m 
f 

X 

1 

1.08 

1.08 







— 

19.  B. 

barrensis 

m 
f 

1 
X 

0.71 

0.71 

— 

— 

— 

— 

20.  B 

cyclodes 

m 

f 

X 

1 

0.80 

0.80 

I 







21.  B 

championi 

m 
f 

X 
X 

— 

— 

— 

— 

— 

— 

22.  B 

ixtatan 

m 
f 

3 
4 

0.58 
0.48 

0.62 
0.60 

0.60 

0.55 

0.01 
0.03 

0.02 
0.05 

3.49 
9.52 

'X"  =  sex  not  available  for  measurement. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


465 


with  arcuate  side  margins,  markedly 
convergent  behind  and  abruptly  sin- 
uate just  before  hind  angles  

18.  B.  armuelles,  new  species 

Pronotum  (Fig.  18)  moderately  con- 
vex, side  margins  not  markedly  ar- 
cuate nor  much  convergent  behind, 
shallowly  sinuate  before  hind  angle 

16.  B.  cortes,  new  species 

12(8).     Elytra  with  pale  spots  contrasting 

against  dark  background  13 

Elytra  concolorous 14 

13(12).  Pronotum  (Fig.  5)  broad,  wider  than 
head  across  eyes;  elytral  pale  spot  ex- 
tended to  apex  in  most  specimens;  in- 
terneur  punctures  coarsely  impressed 
and  large;  pronotum  with  fine  and  fee- 
ble carina  at  hind  angle 

3.  B.  mexicanum  Dejean 

Pronotum  (Fig.  9)  narrow,  about  as 
wide  as  head  across  eyes;  elytral  pale 
spot  isolated;  interneur  punctulae 
finely  impressed,  medium-sized; 
pronotum  with  well-developed  carina 
delimiting  deep  basal  fovea  

7.  B.  quetzal,  new  species 

14(12).  Elytral  interneurs  coarsely  punctate 
basally,  absent  behind  middle,  dorsal 
surface  without  microsculpture,  shiny 

1.  B.  purulha,  new  species 

Elytral  interneurs  finely  punctate  or 
not,  extended  to  near  apex;  dorsal 
surface  with  or  without  microsculp- 
ture, if  without,  hind  wings  absent 

15 

15(14).  Pronotum  (Fig.  6)  with  short,  sharp, 
well-developed  carina  at  hind  angle; 
dorsal  surface  with  metallic-blue  cast; 

flight  wings  fully  developed  

4.  B.  vernale  Bates 

Pronotum  with  rudimentary  or  no  ca- 
rina at  hind  angle;  dorsal  surface  non- 
metallic  brown  or  black;  flight  wings 
reduced  or  virtually  absent  16 

16(15).  Elytron  with  interneurs  7  and  8  vir- 
tually absent  (a  few  minute  punctulae 
present  in  some  individuals);  small 
beetles  with  narrow  ovoid  elytra  and 

narrow  pronotum 

II.  B.  aeger,  new  species 


Elytron  with  all  interneurs  present, 
though  finer  laterally,  all  easily  trace- 
able beyond  disc;  larger  beetles,  with 
broadly  ovoid  elytra  and  broad  prono- 
tum   17 

17(16).  Form  short,  broad,  and  subconvex; 
pronotum  abruptly  sinuate,  sides 
straight  for  only  a  short  distance,  hind 

angles  acute  or  square  18 

Form  long,  narrow,  and  depressed; 
pronotum  with  lateral  margins  nearly 
straight  in  basal  fifth,  hind  angles 
acute  21 

18(17).  Metasternum  with  intercoxal  process 
narrowly  margined,  bead  well  devel- 
oped and  of  even  width 

8.  B.  diabola,  new  species 

Metasternum  with  intercoxal  process 
broadly  margined,  bead  widely  re- 
moved from  apex  medially,  narrowed 
laterally  19 

19(18).  Eyes  flattish,  not  produced  much  be- 
yond outline  of  head  capsule  (range: 

Cerro  Buenavista,  Costa  Rica)  

10.  B.  edwardsi  new  species 

Eyes  prominent,  produced  well  be- 
yond outline  of  head  capsule  20 

20(19).  Range:  central  cordillera  of  Costa 
Rica;  male  aedeagus  as  in  Figure  30 

9.  B.  vulcanium  Darlington 

Range:  Volcan  de  Chiriquf,  Panama; 
male  aedeagus  as  in  Figure  33  

12.  B.  chiriqui,  new  species 

21(17).  Elytron  with  interneurs  markedly 
striatopunctulate,  intervals  subcon- 
vex    5.  B.  satellites  Bates 

Elytron  with  interneurs  shallowly 

striatopunctulate,  intervals  flat  

6.  B.  lavernae,  new  species 

The  semistriatum  group 

(Subgenus  Hydriomicrus  Casey,  1918:87.  Type-species  Bem- 
bidium  semistriatum  Haldeman,  1843:303,  by  subsequent  des- 
ignation of  Lindroth  1963:305.) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered;  elytral  interneurs 
effaced  in  apical  half,  markedly  punctulate  in 
basal  half;  microsculpture  absent;  male  genital 
sclerites  as  in  or  similar  to  Figure  23. 


466 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


FIGURES  3  to  14.  Pronotum,  dorsal  aspect.  3.  Bembidion  purulha  n.sp.,  66.4  km  south  of  Coban,  Alta  Verapaz,  Guatemala, 
male.  4.  B.  rogersi  Bates,  7.5  km  south  of  Ojo  de  Agua,  Cartago,  Costa  Rica,  male.  5.  B.  mexicanum  Dejean,  22.0  km  west 
of  San  Pedro  Sacatepequez  at  Rio  Nahuala,  male.  6.  B.  vernale  Bates,  San  Isidro  de  Coronado,  San  Jose,  Costa  Rica,  male. 
7.  B.  satellites  Bates,  Chomogo  area,  10°18'N,  084°47'W,  Alajuela,  Costa  Rica,  male.  8.  B.  lavernae  n.sp.,  Cerro  Buenavista, 
San  Jose,  Costa  Rica,  female.  9.  B.  quetzal  n.sp.,  20.0  km  west  of  San  Pedro  Sacatepequez  at  Rio  Nahuala,  male.  10.  B. 
diabola  n.sp.,  Volcan  Irazii,  Cartago,  Costa  Rica,  male.  II.  B.  vulcanium  Darlington,  7.5  km  south  of  Ojo  de  Agua,  Cartago, 
Costa  Rica,  male.  12.  B.  edwardsi  n.sp.,  Cerro  Buenavista,  San  Jose,  Costa  Rica,  male.  13.  B.  aeger  n.sp.,  Cerro  Buena- 
vista, San  Jose,  Costa  Rica,  male.  14.  B.  chiriqui  n.sp.,  8.0  km  west  of  Boquete,  Chiriqui,  Panama,  male. 


These  beetles  are  hygrophilous,  occurring 
along  streams  in  fine  gravel  and  sand,  or  in 
Sphagnum  bogs.  Previously,  this  group  of  five 
species  was  known  from  only  North  America; 
the  Central  American  one  described  below  ex- 
tends the  group's  range  considerably.  All 
species  are  fully  winged  and  occur  in  the  low- 
land or  adjacent  low  hills.  The  male  median  lobe 
of  the  species  covered  herein  lacks  the  pig- 
mented  ostium  flag  characteristic  of  Central 
American  Bembidion  (Peryphus)  species,  yet 
the  endophallus  strongly  suggests  relationship 
with  this  group. 

1.  Bembidion  purulha,  new  species 

(Figures  3,  23,  64) 

DIAGNOSTIC  COMBINATION. — Interneurs  with 
large,  coarse,  sparse  punctulae;  pronotum  mark- 
edly constricted  with  small  basal  punctiform 
fovea;  dorsal  surface  without  microsculpture; 
male  median  lobe  without  pigmented  ostium 


flags.  Color  and  luster:  piceous;  appendages  tes- 
taceous; venter  and  dorsum  of  forebody  rufo- 
piceous;  surface  very  shiny.  Form:  medium- 
sized  for  genus  with  narrow  forebody  and  broad 
elytra;  pronotum  (Fig.  3)  slightly  wider  than 
head  or  elytron;  eyes  large  and  prominent. 
Structure:  frontal  furrows  deeply  sulcate  and 
arcuate  between  eyes,  shallower  where  pro- 
longed on  clypeus.  Hind  angles  of  pronotum 
finely  and  abruptly  carinate,  carinae  very  short; 
basal  fovea  deep,  extended  to  lateral  margin. 
Interneurs  effaced  apically;  except  first  which  is 
striate  in  apical  third,  otherwise  coarsely  punc- 
tulate,  punctures  deep  and  large,  separated  by 
their  own  diameter  or  more.  Dorsal  microsculp- 
ture absent.  Male  median  lobe  as  in  Figure  23. 
Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  64. 
Known  only  from  the  Sierra  de  las  Minas  in  east- 
central  Guatemala.  I  have  seen  specimens  from 
localities  listed  in  Table  5. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


467 


16 


1.0mm 


FIGURES  15  to  22.  Pronotum,  dorsal  aspect.  15.  Bembidion  nahuala  n.sp.,  27.0  km  northwest  of  San  Marcos,  San  Marcos, 
Guatemala,  male.  16.  B.  franiae  n.sp.,  4.8  km  east  of  San  Mateo  Ixtatan,  Huehuetenango,  Guatemala,  male.  17.  B.  aratum 
LeConte,  La  Lima,  Cortes,  Honduras,  male.  18.  B.  cortes  n.sp.,  La  Lima,  Cortes,  Honduras,  male.  19.  B.  sparsum  Bates, 
Tikal,  Del  Peten,  Guatemala,  male.  20.  B.  armuelles,  n.sp.,  Puerte  Armuelles,  Chiriqui,  Panama,  female.  21.  B.  championi 
Bates,  8.6  miles  [13.8  km]  east  of  San  Cristobal,  Chiapas,  Mexico,  female.  22a.  B.  ixtatan  n.sp.,  27.0  miles  [43.5  km]  west  of 
Solola,  Solola,  Guatemala,  male.  22b.  B.  barrensis  n.sp.,  Barro  Colorado  Island,  Panama,  male. 


NATURAL  HISTORY  NOTES. — These  beetles 
were  collected  in  May  and  August  at  elevations 
of  1372  m  and  1420  m  in  Lower  Montane  Wet 
and  Subtropical  Wet  forests,  respectively;  nei- 
ther is  teneral.  The  holotype  was  collected  from 
oak  leaf  litter  near  a  stream;  the  paratype  was 
collected  by  splashing  fine  silty  gravel  at  the 
edge  of  a  small  brook  in  pine-oak  forest.  Both 
specimens  are  fully  winged. 

TYPE-MATERIAL. — Holotype  9 ,  GUATEMALA,  Baja  Vera- 
paz,  22.0  km  S  of  Purulha,  15°07'N,  090°12'W,  Aug.  (Ball, 
Frania,  &  Whitehead)  (USNM),  ADP  026375.  Paratype:  Id, 
GUATEMALA,  Aha  Verapaz,  66.4  km  S  Coban,  15°12'N, 
090°18'W,  May  (Erwin  &  Erwin)  (USNM),  ADP  013626.  See 
also  Table  5. 

DERIVATION  OF  TAXON  NAME. — Spanish, 
Purulha,  after  the  town  near  which  the  holotype 
was  collected. 

The  rogersi  group 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  without  border;  elytral  interval  3  bifo- 
veate,  fovea  broad  and  depressed;  male  median 
lobe  with  broad  lobe  on  ventral  face. 

These  beetles  are  independent  of  running  or 
standing  water  and  occur  in  Subparamo  grass- 
lands of  volcanic  highlands.  Two  Colombian 
species  belong  to  this  group:  B.  subapterum 
Darlington  and  B.  sanctaemarthae  Darlington. 
The  three  species  are  either  wing-dimorphic  or 


brachypterous  and  form  a  group  closely  related 
to  the  vernale  group.  The  vernale  group  is 
northern  and  islandic  in  distribution  while  the 
rogersi  group  is  southern;  B.  rogersi  is  the 
northernmost  representative  of  the  group. 

2.  Bembidion  rogersi  Bates 

(Figures  2,  4,  24,  40,  64) 

Bembidium  rogersi  BATES,  1878:602.  [Lectotype  6,  COSTA 
RICA,  Volcan  de  Irazii,  09°58'N,  083°53'W  (BMNH),  here 
designated.] 

DIAGNOSTIC  COMBINATION. — Interneur  3  cat- 
enate, constrictions  at  large  unisetose  foveae; 
color  black.  Color  and  luster:  black;  appendages 
and  venter  piceous;  surface  of  elytron  dull, 
forebody  shiny.  Form:  medium-sized  for  genus 
with  narrow  forebody  and  elytra;  elytra  either 
with  square  humeri  or  sloped;  pronotum  (Fig.  4) 


TABLE  5.    LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  01.  Bembidion  purulha  n.sp.;  map  Figure  64. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month          No. 
coll.           spec. 

15°07'N 
90°12'W 

1420 

USNM 

Aug                   1 

15°12'N 
90°18'W 

1372 

USNM 

May                  1 

Total  specimens  examined:      2 

468 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


TABLE  6.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  02.  Bembidion  rogersi  n.sp.;  map  Figure  64. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month 
coll. 

No. 
spec. 

10°11'N 
84°13'W 

2600 

USNM 

Mar 

1 

09°59'N 

83°53'W 

3000-3030 

USNM 

1 

1 

09°58'N 

83°53'W 

3364 

UASM 

Aug 

6 

09°58'N 

83°53'W 

3440 

MCZ 

Jan 

2 

09°58'N 
83°53'W 

3350 

USNM 

May 

9 

09°55'N 
84°02'W 

1400-1500 

USNM 

Feb 

2 

09°54'N 
83°41'W 

0640 

USNM 

May 

1 

09°54'N 
83°41'W 

0640 

MCZ 

Jun,  Jul 

3 

09°35'N 
83°48'W 

2682 

USNM 

Jun 

182 

09°33'N 

83°44'W 

2700 

UAIC 

Jul 

1 

09°32'N 
83°46'W 

? 

MCZ 

Dec 

2 

09°32'N 

83°46'W 

1 

UAIC 

Mar 

1 

08°51'N 
82°33'W 

2240 

USNM 

Jun 

4 

08°51'N 
82°34'W 

2134 

UAIC 

May 

1 

08°51'N 
82°34'W 

2134-2438 

UAIC 

May 

5 

08°51'N 
82°36'W 

1848-2464 

USNM 

May 

5 

08°47'N 
83°30'W 

2220 

USNM 

Jun 

1 

Total  specimens  examined: 

227 

wider  than  head,  about  equal  in  width  to  elytron; 
eyes  moderately  large  and  prominent.  Struc- 
ture: frontal  furrows  moderately  well  impressed, 
diffuse,  rugose,  parallel,  prolonged  on  clypeus; 
anterior  supraorbital  pore  well  impressed.  Hind 
angles  of  pronotum  markedly  carinate;  basal 
fovea  deep,  rather  small,  extended  to  base;  side 
slightly  sinuate.  Elytron  (Figs.  2a,  b)  either  long, 
humerus  squared,  sides  slightly  arcuate,  or 
short,  humerus  rounded/sloped,  sides  markedly 
arcuate;  interneurs  entire,  much  shallower  api- 
colaterally  and  striate;  interneurs  2  and  3  cate- 
nate at  Ed3a  and  Ed5b.  Dorsal  microsculpture 
(Fig.  40)  of  rather  large,  coarse,  transverse 


meshes,  effaced  from  center  of  pronotum.  Male 
median  lobe  as  in  Figure  24.  Size:  see  Tables  2, 
3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  64; 
known  from  the  Cordillera  Central  of  Costa  Rica 
from  Volcan  Irazii,  Volcan  Poas,  and  in  the  area 
of  Turrialba,  south  along  the  Cordillera  de  Ta- 
lamanca  to  Volcan  de  Chiriqui  in  Panama. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  commonly  collected  above  1600  m  in  life 
zones  from  Lower  Montane  Wet  Forest  up  and 
into  Subparamo.  They  occur  as  high  as  3350  m; 
are  found  independent  of  running  water;  and 
have  been  collected  in  January,  February,  March, 
May  through  August,  and  December.  Very  tener- 
al  beetles  were  found  in  late  May  on  Volcan  de 
Chiriqui,  indicating  that  larvae  and  pupae  may 
be  found  just  prior  to  that  time,  that  is,  in  the 
early  rainy  season.  Great  numbers  of  specimens 
were  collected  at  night  on  fine-textured  wet  soil 
among  bunch  grass  at  the  edge  of  dwarf  Mon- 
tane Rain  Forest  on  Cerro  de  la  Muerte,  Costa 
Rica  (2682  m),  and  among  flood  debris  and  under 
stones  during  the  day  near  Cerro  Punta,  Chiri- 
qui, Panama.  Individuals  of  this  species  may  be 
long- winged  and  presumably  fly,  or  brachypter- 
ous,  having  only  a  small  veinless  pad  in  place  of 
the  flight  wing.  Table  1  illustrates  the  distribu- 
tion of  these  character  states  at  several  locali- 
ties. 

MATERIAL  EXAMINED. — Lectotype  (see  above)  and  227 
specimens  (Table  6). 

The  vernale  group 

(Subgenus  Peryphus  Stephens,  1828:2.  [Type-species  Cara- 
bus  litorale  auct.  =  Bembidium  tetracolum  Say,  1823:89,  by 
subsequent  designation  of  Westwood  1840:7.] 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered;  elytron  with  dis- 
cal  setae  contiguous  with  or  in  interneur  3 ;  par- 
ameres  trisetose  or  bisetose;  endophallus  as  in 
Figures  25  to  33;  elytral  microsculpture  of  trans- 
verse lines  or  meshes,  effaced  from  head  and 
pronotum;  interneurs  shallow  or  effaced  later- 
ally and  apically. 

A  study  of  the  life  histories  of  these  beetles 
would  add  much  to  test  the  theory  of  taxon 
pulses  (Erwin  1979,  198  la).  It  is  readily  apparent 
that  the  group  is  plesiotypically  hygrophilous. 
With  development  of  aptery  and  movement  to 
montane  habitats,  there  is  a  corresponding  shift 
away  from  water  into  forest-floor  litter  or  open 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


469 


dry  spots  in  the  forest  or  above  treeline,  thus 
the  species  become  mesophilous  alticoles.  In 
Central  America,  this  shift  was  accomplished  by 
three  separate  lineages  of  the  vernale  group. 
The  plesiotypic  parent  stock  was  probably  a 
middle  altitude  riparian  species  which  radiated 
and  is  now  represented  by  four  widespread 
species  at  middle  altitudes  one  of  which  is  wing- 
dimorphic  and  six  brachypterous  alticoles. 

As  suggested  by  Lindroth  (1963:312),  a  world- 
wide revision  of  this  group  would  be  desirable. 
Only  then  will  we  know  the  subgroup  relation- 
ships, and  these  can  be  recognized  only  by  the 
structure  of  the  endophallus  in  combination  with 
other  characters.  The  median  lobe  of  all  species 
in  this  group  has  one  or  two  ostium  flags  ex- 
tending from  near  the  apical  orifice  to  the  me- 
dian area  on  the  left  surface.  In  addition,  three 
Central  American  forms  have  a  nonsclerotized 
patch  ventrally  on  the  median  lobe.  These  apo- 
typic  features  should  assist  in  discovering  rela- 
tionships in  revisions  with  wider  geographic 
coverage  (e.g.,  Erwin  and  Kavanaugh  1981). 

Widespread  species  exhibit  considerable  vari- 
ation in  pronotal  shape  and  elytral  coloration. 
Bembidion  mexicanum  and  B.  vernale  exempli- 
fy this  in  part  in  regard  to  pronotal  variation, 
and  the  former  is  paler  in  the  north,  darker  in 
the  south. 


3.  Bembidion  mexicanum  Dejean 

(Figures  5,  25,41,65) 

Bembidium  mexicanum  DEJEAN,  1831:126.  [Lectotype  d, 
MEXICO  (MNHP)  selected  and  so  labelled  by  G.  E.  Ball.] 

Bembidium  sallaei  BATES,  1882:148.  [Lectotype  3,  MEXICO 
(MNHP)  selected  and  so  labelled  by  G.  E.  Ball.] 

DIAGNOSTIC  COMBINATION. — Pronotum  (Fig. 
5)  broad,  wider  than  head  across  eyes;  elytral 
pale  spot  extended  to  apex  in  most  specimens; 
interneur  punctures  coarsely  impressed  and 
large;  pronotum  with  fine  and  feeble  carina  at 
hind  angle.  Color  and  luster:  dorsum  piceous; 
appendages  and  at  least  part  of  elytron  testa- 
ceous; venter  piceous  anteriorly,  abdomen  ru- 
fous posteriorly;  surfaces  shiny,  elytron  slightly 
iridescent.  Form:  large  beetles  with  broad  head 
and  pronotum,  and  broad  elytra;  pronotum  (Fig. 
5)  wider  than  head,  about  equal  to  width  of  ely- 
tron; eyes  large  and  prominent.  Structure:  fron- 
tal furrows  well  impressed,  bowed  medially 
around  frontal  callus,  prolonged  on  clypeus, 
generally  parallel.  Hind  angle  of  pronotum  finely 


carinate,  carina  curved  posteriolaterally;  basal 
fovea  well  impressed,  narrowed  anteriorly.  In- 
terneurs  effaced  apicolaterally,  7  virtually  ab- 
sent, 1  to  3  striatopunctulate,  4  to  6  punctulate. 
Dorsal  microsculpture  of  extremely  fine  and 
dense  transverse  lines  on  elytron  (Fig.  41),  ef- 
faced from  head  and  pronotum.  Male  median 
lobe  as  in  Figure  25.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  65. 
At  present,  this  species  is  known  from  Colorado 
south  to  middle  Costa  Rica. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  commonly  found  at  middle  elevations  in 
Lower  Montane  Wet,  Premontane  Moist,  Trop- 
ical Dry,  and  Subtropical  Dry  forests  in  the  pine 
zone  between  1300  m  and  2400  m.  Specimens 
were  collected  in  March,  May,  June,  and  Au- 
gust; a  teneral  specimen  from  Guatemala  col- 
lected in  May  indicates  that  larvae  and  pupae 
may  be  found  just  prior  to  that  time,  that  is,  in 
the  early  rainy  season.  They  are  found  at  the 
edge  of  small  to  medium-sized  streams  in  gravel 
and  under  stones  where  there  is  some  silt  and 
at  least  some  sparse  vegetation.  All  known  spec- 
imens are  fully  winged;  P.  J.  Spangler  collected 
two  individuals  at  UV  light  traps  near  Guate- 
mala City  indicating  that  the  beetles  fly. 

MATERIAL  EXAMINED. — Lectotypes  (see  above)  and  62 
specimens  (Table  7). 

TAXONOMIC  NOTES. — Bates' s  B.  sallaei  is  a 
color  variety  of  the  widespread  and  variable  B. 
mexicanum,  and  was  recognized  as  such  by 
Bates  (1882: 148). 

4.  Bembidion  vernale  Bates 

(Figures  1,6,  26,42,66) 

Bembidium  vernale  BATES,  1882:149.  [Lectotype  d  ,  PANAMA, 
Pena  Blanca,  08°27'N,  081041'W  (BMNH),  here  designat- 
ed.] 

DIAGNOSTIC  COMBINATION. — Elytra  blue, 
nonspotted;  length  more  than  4.2  mm.  Color 
and  luster:  dorsum  submetallic  blue;  penulti- 
mate palpomeres  and  antennomeres  2  to  1 1  in- 
fuscated,  otherwise  appendages  testaceous; 
venter  piceous;  surface  shiny.  Form:  large-sized 
beetles  for  genus  with  narrow  forebody  and 
slightly  broad  elytra.  Pronotum  (Fig.  6)  slightly 
wider  than  head,  about  equal  to  width  of  elytron; 
eyes  large  and  prominent.  Structure:  frontal  fur- 
rows well  impressed,  rugose,  slightly  convergent 
anteriorly,  extended  on  clypeus.  Hind  angle  of 


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35 


38 


39b 


FIGURES  23  to  39.  Median  lobe  of  male  genitalia,  left  lateral  aspect.  23.  Bembidion  purulha  n.sp.,  66.4  km  south  of  Coban, 
Alta  Verapaz,  Guatemala.  24.  B.  rogersi  Bates,  7.5  km  south  of  Ojo  de  Agua,  Cartago,  Costa  Rica.  25.  B.  mexicanum  Dejean, 
22.0  km  west  of  San  Pedro  Sacatepequez  at  Rio  Nahuala.  26.  B.  vernale  Bates,  San  Isidro  de  Coronado,  San  Jose,  Costa  Rica. 
27.  B.  satellites  Bates,  Chomogo  area,  10°18'N,  084°47'W,  Alajuela,  Costa  Rica.  28.  B.  quetzal  n.sp.,  25.0  km  south  of  Toto- 
nicapan,  Huehuetenango,  Guatemala.  29.  B.  diabola  n.sp.,  Volcan  Irazii,  Cartago,  Costa  Rica.  30.  B.  vulcanium  Darlington, 
7.5  km  south  of  Ojo  de  Agua,  Cartago,  Costa  Rica.  31.  B.  edwardsi  n.sp.,  Cerro  Buenavista,  San  Jose,  Costa  Rica.  32.  B. 
aeger  n.sp.,  Cerro  Buenavista,  San  Jose,  Costa  Rica.  33.  B.  chiriqui  n.sp.,  8.0  km  west  Boquete,  Chiriqui,  Panama.  34.  B. 
nahuala  n.sp.,  27.0  km  northwest  of  San  Marcos,  San  Marcos,  Guatemala.  35.  B.  franiae  n.sp.,  4.8  km  east  of  San  Mateo 
Ixtatan,  Huehuetenango,  Guatemala.  36.  B.  aratum  LeConte,  La  Lima,  Cortes,  Honduras.  37.  B.  cortes  n.sp.,  La  Lima, 
Cortes,  Honduras.  38.  B.  sparsum  Bates,  Tikal,  Del  Peten,  Guatemala.  39a.  B.  ixtatan  n.sp.,  27.0  miles  [43.5  km]  west  of 
Solola,  Solola,  Guatemala.  7>9b.  B.  barrensis  n.sp.,  Barro  Colorado  Island,  Panama. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


471 


pronotum  markedly  carinate;  basal  fovea  large 
and  deep.  Interneurs  entire,  shallow  laterally 
and  apically,  punctulate  throughout  length.  Dor- 
sal microsculpture  of  extremely  fine  and  dense 
transverse  lines  or  meshes  (more  on  female  than 
male)  on  elytron  (Fig.  42),  effaced  from  head 
and  pronotum.  Male  medium  lobe  as  in  Figure 
26.  Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  66; 
the  range  of  this  species  extends  in  the  north 
from  at  least  Guatemala,  south  to  western  Pan- 
ama; no  specimens  from  Mexico  were  exam- 
ined. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  in  large  numbers  at  middle  altitudes 
along  rivers  with  stony  and  gravelly  banks 
where  there  is  little  or  no  vegetation.  Altitudinal 
range  extends  from  1700  m  to  2600  m  in  Pre- 
montane  Wet,  Lower  Montane  Moist,  Lower 
Montane  Wet,  and  Montane  Wet  forests.  Spec- 
imens were  collected  in  April,  May,  and  June; 
teneral  ones  collected  in  May  indicate  that  lar- 
vae and  pupae  may  be  found  just  prior  to  that 
time,  that  is,  in  the  early  rainy  season.  Individ- 
uals occur  at  the  edge  of  the  water,  hiding  under 
stones  during  the  day.  All  individuals  studied 
were  fully  winged  and  numerous  collection  rec- 
ords indicate  capture  at  UV  lights;  these  beetles 
are  probably  good  fliers. 

MATERIAL  EXAMINED. — Lectotype  (see  above)  and  295 
specimens  (Table  8). 

TAXONOMIC  NOTES. — Bates,  in  his  original 
description,  gave  as  type-locality  of  this  species, 
"Mexico  (Salle),"  however,  he  specifically 
mentioned  later  (Bates  1884:291)  that  additional 
material  from  Pena  Blanca,  Panama,  was  iden- 
tifiable as  B.  vernale.  The  type-specimen  from 
the  Salle  collection  is  apparently  lost,  therefore, 
I  selected  one  of  the  Pena  Blanca  specimens  as 
lectotype,  thereby  restricting  the  type-locality. 

5.  Bembidion  satellites  Bates 

(Figures  7,  27,  43,  67) 

Bembidium  satellites  BATES,  1884:291.  [Lectotype  9,  PANA- 
MA, Pena  Blanca,  08°27'N,  08r41'W  (BMNH),  here  des- 
ignated.] 

DIAGNOSTIC  COMBINATION. — Form  long, 
narrow,  and  depressed;  pronotum  with  lateral 
margins  nearly  straight  in  basal  fifth,  hind  angles 
acute.  Elytron  with  interneurs  markedly  stria- 
topunctulate,  intervals  subconvex.  Dorsal  sur- 


TABLE  7. 
LABELS.  03. 

LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
Bembidion  mexicanum  Dejean;  map  Figure  65. 

Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month 
coll. 

No. 
spec. 

15°20'N 
91°26'W 

ca.  1900 

USNM 

Jun 

5 

15°12'N 
90°18'W 

1372 

USNM 

May 

20 

15°07'N 
91°32'W 

ca.  2000 

USNM 

Jun 

2 

15°02'N 
90°27'W 

1400 

USNM 

Jun 

1 

14°58'N 
91°46'W 

2200 

USNM 

May 

24 

14°49'N 
91°02'W 

2370 

USNM 

Aug 

1 

14°37'N 
90°30'W 

1479 

MCZ, 

UAIC 

9 

3 

14°37'N 
90°30'W 

1479 

USNM 

? 

1 

14°37'N 
90°30'W 

1525 

USNM 

9 

1 

14°33'N 
90°35'W 

ca.  1300 

USNM 

Aug 

2 

09°55'N 
84°02'W 

1000-1200 

USNM 

Feb-Mar 

1 

09°55'N 
84°02'W 

1000-1200 

USNM 

Jun 

1 

Total  specimens  examined: 

62 

face  without  metallic-blue  cast.  Color  and  lus- 
ter: dorsum  flavous;  antennomeres  3  to  11 
infuscated,  otherwise  appendages  testaceous; 
venter  piceous;  surface  shiny,  elytra  slightly  ir- 
idescent. Form:  medium-sized  beetles  with  nar- 
row forebody  and  very  narrow,  depressed  ely- 
tra; pronotum  (Fig.  7)  convex,  slightly  wider 
than  head  and  elytron;  eyes  large  and  promi- 
nent. Structure:  frontal  furrows  well  impressed, 
arcuate  medially,  extended  on  clypeus.  Hind  an- 
gle of  pronotum  markedly  carinate;  basal  fovea 
large  and  deep.  Interneurs  effaced  apicolateral- 
ly,  or  nearly  so;  interneur  7  with  small  number 
of  widely  spaced  minute  punctulae,  1  to  6  stria- 
topunctulate.  Dorsal  microsculpture  of  ex- 
tremely fine  and  dense  transverse  lines  or  mesh- 
es on  elytron  (Fig.  43),  effaced  from  head  and 
pronotum.  Male  median  lobe  as  in  Figure  27. 
Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  67; 
the  range  of  this  species  extends  from  Monte- 


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TABLE  8. 

LOCALITY 

DATA  DEDUCED  FROM  SPECIMEN 

TABLE  9.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 

LABELS.  04.  Bembidion  vernale  Bates; 

map  Figure  66. 

LABELS.  05.  Bembidion  satellites  Bates;  map  Figure  67. 

Elev. 

Orig. 

Month 

No. 

Elev.                 Orig.           Month         No. 

Lat./Long. 

(m) 

deposit. 

coll. 

spec. 

Lat./Long.            (m)                deposit.          coll.          spec. 

15°15'N 

10°18'N 

91°27'W 

2560 

USNM 

Aug 

1 

1300-1600            USNM           Mar               4 
84  48  W 

10°15'N 

10°18'N 

0010 

USNM 

Jun 

1 

1620                 USNM           Jun                7 

83°28'W 

84°47'W 

09°59'N 

08°5  1  'N 

84°00'W 

1524 

UASM 

Jun 

13 

1848-2464            USNM           May             32 
82  35  W 

08°51'N 

08°51'N 

82°36'W 

2240 

USNM 

Jun 

4 

o<>o,c,«,           1848-2464            USNM           May               5 
82  35  W 

08°51'N 

08°51'N 

82°36'W 

1848-2464 

USNM 

May 

112 

82'34'W           2134-2438           UAIC            May              6 

08°51'N 
82°36'W 

1720 

UASM 

May 

14 

2134-2438            USNM           May              2 
82  34  W 

08°51'N 

1720 

HPST 

Apr,  May 

9 

Total  specimens  examined:     56 

08°51'N 

1700 

USNM 

Jun 

35 

82°36'W 

08°51'N 

2134-2438 

UAIC 

May 

5 

82°34'W 

membrane.  Within  populations  the  length  of  the 

08°51'N 

Q*)O1A  '\\7 

2134-2438 

UAIC 

May 

13 

wing  membrane  varied  from  a  small  veined  pad 

82  34  W 

to  a  longer  veined  membrane  one-third  the 

08°51'N 
82°34'W 

2134-2438 

USNM 

May 

12 

length  of  the  elytron,  to  a  fully  developed  wing 

with  reflexed  apex.  Such  variation  was  noted 

08°47'N 

82°26'W 

1848 

UAIC 

Jun 

5 

from  both  Monteverde,  Costa  Rica,  and  Chiri- 

08°47'N 

qui,  Panama. 

1848 

USNM 

Jun 

69 

82°26'W 

MATERIAL   EXAMINED.  —  Lectotype   (see   above)   and   56 

08°47'N 

1100 

USNM 

Jun 

1 

specimens  (Table  9). 

82°26'W 

08°46'N 

1360 

HPST 

May 

1 

6.  Bembidion  lavernae,  new  species 

82°38'W 

(Figures  8,  67) 

Total  specimens  examined: 

295 

DIAGNOSTIC  COMBINATION.  —  Elytral   inter- 

verde,  Costa  Rica,  south  along  the  Cordillera  de 
Talamanca  to  Volcan  de  Chiriquf,  Panama. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  middle  altitudes  along  slow,  small 
streams  with  silty  and  gravelly  banks  where 
there  is  little  or  no  vegetation.  Altitudinal  range 
extends  from  1600  m  to  2500  m  in  Premontane 
Rain  and  Lower  Montane  Wet  forests.  Speci- 
mens were  collected  in  March,  May,  and  June; 
teneral  ones  collected  in  May  and  mid-June  at 
Chiriqui  indicate  that  larvae  and  pupae  may  be 
found  just  prior  to  that  time,  that  is,  in  the  early 
rainy  season.  Individuals  occur  at  the  edge  of 
the  water,  hiding  among  small  stones  during  the 
day.  Individuals  studied  were  either  fully  winged 
or  with  some  state  of  reduction  of  the  wing 


vals  flat;  elytra  elongate  and  spatulate;  prono- 
tum  with  lateral  margins  nearly  straight  in  basal 
fifth,  hind  angles  acute.  Color  and  luster:  rufo- 
piceous;  head,  prothorax,  venter  flavous,  in  part 
infuscated;  appendages  flavotestaceous,  in  part 
infuscated;  surfaces  shiny.  Form:  medium-sized 
for  genus  with  head  and  prothorax  robust;  elytra 
long  and  spatulate,  with  markedly  sloped  humeri 
and  markedly  arcuate  sides;  pronotum  (Fig.  8) 
markedly  narrowed  basally,  slightly  wider  than 
head,  much  wider  than  elytron;  eyes  medium- 
sized,  slightly  prominent.  Structure:  frontal  fur- 
rows well  impressed,  rugose,  parallel,  ended 
abruptly  at  frontoclypeal  line;  clypeus  convex. 
Hind  angle  of  pronotum  noncarinate,  square; 
fovea  broad  and  deep;  side  margins  straight  in 
basal  fifth.  Interneurs  1  and  2  entire,  striate,  3 
to  6  abbreviated  before  apex,  7  and  8  effaced. 
Dorsal  microsculpture  of  moderately  transverse 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


473 


meshes.  Male  unknown.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  67; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — These  beetles 
live  microsympatrically  with  B.  edwardsi  on 
Cerro  Buenavista  at  high  elevation  (3416  m)  in 
the  Subparamo  plant  association  (Janzen  1973), 
Montane  Rain  Forest  of  Holdridge  (1971).  The 
type  is  brachypterous,  as  are  no  doubt  the  rest 
of  the  members  of  the  species  since  humeral  re- 
duction is  more  marked  than  in  any  other 
species  studied  except  B.  aeger. 

TYPE-MATERIAL. — Holotype  $ ,  COSTA  RICA,  San  Jose 
Province,  Cerro  Buenavista,  09°33'N,  083°45'W,  Mar.  (Ed- 
wards) (USNM),  ADP  055189. 

DERIVATION  OF  TAXON  NAME. — I  take  great 
pleasure  in  naming  this  species  in  honor  of  La 
Verne  J.  Magadan  who  trudged  up  many  moun- 
tains in  search  of  carabid  beetles,  and  who  de- 
signed the  computerized  system  employed  in 
this  and  my  other  carabid  studies. 


GEOGRAPHICAL  DISTRIBUTION. — Figure  67; 
known  only  from  west-central  Guatemala. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  commonly  at  middle  altitudes  along 
small  rivers  with  stony  and  gravelly  banks  where 
there  is  little  or  no  vegetation.  Altitudinal  range 
extends  from  2200  m  to  2900  m  in  Lower  Mon- 
tane Wet  and  Montane  Moist  forests.  Specimens 
were  collected  in  May  and  August,  none  were 
teneral.  Individuals  occur  at  the  edge  of  water, 
hiding  under  stones  during  the  day.  All  individ- 
uals studied  were  fully  winged. 

TYPE-MATERIAL.— Holotype  <J ,  GUATEMALA,  Quezalte- 
nango,  20.0  km  w  San  Pedro  Sacatepequez  at  Rio  Nahuala 
and  Hwy  1,  14°48'N,  091°46'W,  May  (Erwin  &  Erwin) 
(USNM),  ADP  012345.  Paratypes:  8cJ ,  182 ,  same  data  as  ho- 
lotype  or  as  listed  in  Table  10  and  deposited  as  USNM- 12, 
UASM-2,  CAS-2,  MCZ-2,  AMNH-2,  BMNH-2,  MNHP-2, 
JNEG-2. 

DERIVATION  OF  TAXON  NAME. — Mayan, 
Quetzal,  after  the  beautiful  Guatemalan  bird, 
Pharomachrus  mocinno,  with  which  these  bee- 
tles share  the  forest. 


7.  Bembidion  quetzal,  new  species 

(Figures  9,  28,  44,  67) 

DIAGNOSTIC  COMBINATION. — Pronotum  nar- 
row, about  as  wide  as  head  across  eyes;  elytral 
pale  spot  preapical  and  isolated;  interneur  punc- 
tulae  finely  impressed,  medium-sized;  hind  angle 
of  pronotum  with  well-developed  straight  carina 
delimiting  deep  basal  fovea;  antennomeres  2  to 
1 1  darkly  infuscated.  Color  and  luster:  dorsum 
piceous;  antennomeres  2  to  11,  palpomeres,  tib- 
iae, and  tarsomeres  infuscated,  otherwise  tes- 
taceous; venter  rufopiceous;  surfaces  shiny. 
Form:  moderately  large  beetles  with  narrow 
head  and  pronotum  and  rather  long,  almost  par- 
allel-sided elytra;  pronotum  (Fig.  9)  slightly  wid- 
er than  head,  about  equal  to  width  of  elytron; 
eyes  large  and  prominent.  Structure:  frontal  fur- 
rows well  impressed,  rugose,  almost  parallel, 
and  extended  on  clypeus.  Hind  angle  of  prono- 
tum markedly  carinate;  basal  fovea  large  and 
deep.  Interneurs  effaced  apicolaterally  or  nearly 
so;  interneur  7  with  small  number  of  widely 
spaced,  minute  punctulae,  1  to  6  striatopunctu- 
late,  at  least  basally.  Dorsal  microsculpture  of 
extremely  fine  and  dense  transverse  lines  or 
meshes  on  elytron  (Fig.  44),  effaced  from  head 
and  pronotum.  Male  median  lobe  as  in  Figure 
28.  Size:  see  Tables  2,  3,  and  4. 


8.  Bembidion  diabola,  new  species 

(Figures  10,  29,  45,  68) 

DIAGNOSTIC  COMBINATION. — Form  short, 
broad,  and  subconvex;  eyes  small  and  flattish; 
pronotum  abruptly  sinuate,  sides  straight  for 
only  a  short  distance,  hind  angle  acute  or 
square,  carina  feebly  developed;  metasternum 
with  intercoxal  process  narrowly  margined, 
bead  well  developed  and  of  even  width.  Color 
and  luster:  dorsum  rufopiceous;  scape  testa- 
ceous, otherwise  appendages  partially  infuscat- 
ed; venter  rufopiceous;  surfaces  shiny.  Form: 
small,  rather  robust  beetles  with  large  head, 
small  eyes,  markedly  cordate  pronotum,  ovate 
elytra;  pronotum  (Fig.  10)  much  wider  than  head 
or  elytron;  eyes  flattish.  Structure:  frontal  fur- 
rows well  impressed,  slightly  rugose,  almost 
parallel,  angularly  prolonged  on  clypeus.  Hind 
angle  of  pronotum  with  feebly  developed  carinae 
or  no  carinae;  basal  fovea  large  and  deep;  side 
margins  abruptly  and  markedly  sinuate.  Inter- 
neurs entire,  though  much  shallower  apicolater- 
ally, striatopunctulate,  punctulae  small  and 
sparse.  Dorsal  microsculpture  of  large  trans- 
verse meshes  on  elytra  (Fig.  45),  effaced  from 
head  and  pronotum.  Male  median  lobe  as  in  Fig- 
ure 29.  Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  68; 


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TABLE  10.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  07.  Bembidion  quetzal  n.sp.;  map  Figure  68. 


TABLE  11.    LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  08.  Bembidion  diabola  n.sp.;  map  Figure  68. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month         No. 
coll.          spec. 

Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month 
coll. 

No. 
spec. 

15°07'N 
91°32'W 

ca.  2000 

USNM 

Jun 

2 

09°58'N 
83°53'W 

3440 

MCZ 

Jan 

3 

15°04'N 
91°52'W 

2800-2900 

USNM 

May 

1 

09°58'N 
83°53'W 

3350 

USNM 

May 

37 

14°58'N 
91°46'W 

2200 

USNM 

May 

23 

09°58'N 
83°53'W 

2850 

UMAA 

Feb 

1 

14°49'N 
91°02'W 

2370 

USNM 

Aug 

1 

09°58'N 
83°53'W 

2850 

UASM 

Sep 

3 

Total  specimens  examined: 

27 

09°58'N 
83°53'W 
?  [Costa 
Rica] 

2844 
7 

AMNH 
UAIC 

Nov 

9 

1 

2 

these  beetles  occur 

only  in  the 

Cordillera 

de  la 

?  [Costa 
Rica] 

1 

UMAA 

Jul 

6 

Talamanca,  Costa  Rica. 

NATURAL  HISTORY  NOTES. — These  beetles 
live  microsympatrically  with  B.  vulcanium  on 
Volcan  Irazii  at  high  elevations  in  the  Subpara- 
mo  plant  association  (Janzen  1973),  Montane 
Wet  Forest  of  Holdridge  (1971),  between  2800 
m  and  3500  m.  They  were  collected  in  January, 
May,  September,  and  November;  teneral  spec- 
imens were  found  in  all  those  months  except 
November,  indicating  a  nonseasonal  life  cycle. 
These  beetles  are  found  independent  of  water 
among  bunch  grasses  on  finely  textured  soil.  All 
individuals  studied  have  small  nonveined  pads 
in  place  of  flight  wings. 

TYPE-MATERIAL.— Holotype  <J ,  COSTA  RICA,  Volcan  Irazii, 
09°58'N,  083°53'W,  May  (Cartwright),  ADP  003062.  Para- 
types:  30cJ,  22$  ,  same  data  as  holotype  or  as  listed  in  Table 
11  and  deposited  as  USNM-27,  UASM-6,  CAS-4,  MCZ-3, 
AMNH-3,  BMNH-2,  MNHP-2,  JNEG-2,  UAIC-2,  UMAA-1. 

DERIVATION  OF  TAXON  NAME. — Latin,  dia- 
bolus,  meaning  the  devil,  whose  lair  lay  in  the 
caldera  of  Volcan  Irazii  near  which  these  beetles 
live. 

9.  Bembidion  vulcanium  Darlington 

(Figures  11,  30,46,68) 

Bembidion  vulcanium  DARLINGTON,  1934:157.  [Holotype  6\ 

COSTA  RICA,  Volcan  Irazu,  09°58'N,  083°53'W,  Jan.  (Nev- 

ermann)  (MCZ),  ADP  003145.] 

DIAGNOSTIC  COMBINATION. — Eyes  promi- 
nent, produced  well  beyond  outline  of  head  cap- 
sule; pronotum  robust,  very  broad;  elytral  mi- 
crosculpture  shallowly  engraved,  surface  very 
shiny.  Color  and  luster:  dorsum  rufopiceous  or 
piceous;  scape  and  femora  testaceous,  other- 


Total  specimens  examined:    53 


wise  appendages  infuscated;  venter  rufopiceous 
or  piceous;  surfaces  shiny.  Form:  moderately 
small  beetles  with  broad  head  and  markedly 
broad  pronotum,  elytra  ovoid;  pronotum  (Fig. 
1 1)  much  wider  than  head  or  elytron,  markedly 
convex  and  cordiform;  eyes  large  and  promi- 
nent. Structure:  frontal  furrows  moderately  well 
impressed,  broad,  parallel,  extended  on  clypeus. 
Hind  angle  of  pronotum  feebly  carinate,  carinae 
not  extended  to  base;  basal  fovea  deep,  extend- 
ed to  side  margin;  side  margins  abruptly  and 
markedly  sinuate.  Interneurs  entire,  though 
much  shallower  laterally  and  apically,  almost 
effaced  in  some  individuals,  striatopunctulate 
medially,  punctulate  laterally,  punctulae  rather 
large  and  coarse  on  disc,  widely  separated.  Dor- 
sal microsculpture  of  shallowly  engraved  trans- 
verse meshes  on  elytra  (Fig.  46),  effaced  from 
head  and  pronotum.  Male  median  lobe  as  in  Fig- 
ure 30.  Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  68; 
known  only  from  the  three  high  volcanos  of  Cos- 
ta Rica:  Poas,  Irazii,  and  Cerro  de  la  Muerte. 

NATURAL  HISTORY  NOTES. — These  beetles 
live  microsympatrically  with  B.  diabola  on  Vol- 
can Irazii  and  at  high  elevations  in  the  Subpa- 
ramo  plant  association  (Janzen  1973),  Montane 
Rain  and  Montane  Wet  forests  of  Holdridge 
(1971),  between  2600  m  and  3400  m  on  other 
Costa  Rican  volcanos.  They  were  collected  in 
January,  February,  March,  June,  July,  and  Au- 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


475 


gust;  teneral  specimens  were  not  found.  These 
beetles  are  found  independent  of  water  among 
bunch  grasses  on  finely  textured  soil  under 
stones  and  pieces  of  wood.  All  individuals  stud- 
ied have  small  nonveined  pads  in  place  of  flight 
wings. 

MATERIAL  EXAMINED. — Holotype  (see  above)  and  32  spec- 
imens (Table  12). 

10.  Bembidion  edwardsi,  new  species 

(Figures  12,31,68) 

DIAGNOSTIC  COMBINATION. — Eyes  small  and 
nearly  flat;  metasternal  process  broadly  mar- 
gined; elytra  very  short  and  narrow  in  propor- 
tion to  head  and  pronotum.  Color  and  luster: 
dorsum  of  elytron  rufopiceous,  forebody  pi- 
ceous;  antennomeres  1,2,  and  base  of  3,  legs, 
and  base  of  palpomeres  testaceous;  venter  pi- 
ceous;  surface  shiny.  Form:  medium-sized  for 
genus  with  large  robust  forebody,  small  elytra; 
pronotum  (Fig.  12)  broader  than  head,  much 
broader  than  elytron;  eyes  small,  flattish.  Struc- 
ture: frontal  furrows  well  impressed,  rugose, 
markedly  delimited  laterally  by  convex  carina; 
anterior  supraorbital  seta  in  deep  fovea.  Hind 
angle  of  pronotum  without  carina,  slightly  ob- 
tuse; basal  fovea  and  base  markedly  rugose;  side 
margins  markedly  sinuate  at  basal  third.  Elytron 
short,  with  slightly  sloped  humerus,  and  arcuate 
side  margin.  Interneurs  1  and  2  entire,  3  to  8 
abbreviated  before  apex,  shallower  laterally,  all 
striatopunctulate.  Dorsal  microsculpture  of  ely- 
tron transverse,  not  regularly  formed  of  meshes, 
effaced  from  head  and  pronotum.  Male  median 
lobe  as  in  Figure  31.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  68; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — These  beetles 
live  microsympatrically  with  B.  lavernae  on 
Cerro  Buenavista  at  3416  m  elevation  in  the  Sub- 
paramo  plant  association  (Janzen  1973).  They 
were  collected  in  March.  These  beetles  are 
found  independent  of  water  among  bunch  grass- 
es on  finely  textured  soil  under  stones.  All  in- 
dividuals studied  have  small  nonveined  pads  in 
place  of  flight  wings. 

TYPE-MATERIAL. — Holotype  <J,  COSTA  RICA,  San  Jose 
Province,  Cerro  Buenavista,  09°33'N,  083°45'W,  Mar.  (Ed- 
wards) (USNM),  ADP  055177.  Paratypes:  29,  same  data  as 
holotype  and  deposited  as  CAS-1,  USNM-1. 

DERIVATION  OF  TAXON  NAME. — I  take  great 


TABLE  12.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  09.  Bembidion  vulcanium  Darlington;  map  Figure  68. 


Lat./Long. 


Elev.  Orig.  Month  No. 

(m)  deposit.  coll.  spec. 


10°11'N 
84°13'W 

2400 

UASM 

Aug 

7 

09°58'N 
83°53'W 

2200 

USNM 

Jan-Feb 

1 

09°58'N 
83°53'W 

2800-3000 

USNM 

Jan 

7 

09°58'N 
83°53'W 

2800-3000 

MCZ 

Jan 

4 

09°35'N 
83°48'W 

2682 

USNM 

Jun 

1 

09°33'N 
83°44'W 

2700 

USNM 

Jul 

3 

09°33'N 
83°42'W 

2900-3000 

GRNO 

Aug 

1 

09°32'N 
83°46'W 

9 

UAIC 

Mar 

3 

09°32'N 
83°46'W 

3203 

USNM 

Mar 

1 

09°32'N 
83°46'W 

3355 

USNM 

Jun 

5 

Total  specimens  examined: 

33 

pleasure  in  naming  this  species  in  honor  of  J. 
Gordon  Edwards,  who  first  showed  me  a  ground 
beetle  and  who  collected  the  type-series. 

11.  Bembidion  aeger,  new  species 

(Figures  13,32,47,69) 

DIAGNOSTIC  COMBINATION. — Pronotum  with 
rudimentary  or  no  carina  at  hind  angle;  dorsal 
surface  nonmetallic  brown  or  black;  flight  wings 
reduced  or  virtually  absent;  elytron  with  inter- 
neurs  7  and  8  virtually  absent  (a  few  minute 
punctulae  present  in  some  individuals);  small 
beetles  with  narrow  ovoid  elytra  and  narrow 
pronotum.  Color  and  luster:  dorsum  piceous; 
scape  and  legs  testaceous  or  slightly  infuscated, 
other  appendages  darkly  infuscated;  venter  pi- 
ceous. Form:  small,  rather  cylindrical  beetles 
with  broad  forebody  in  proportion  to  elytra; 
pronotum  (Fig.  13)  slightly  wider  than  head  or 
elytron;  eyes  small  and  slightly  prominent. 
Structure:  frontal  furrows  shallow,  slightly  con- 
vergent, extended  on  clypeus.  Hind  angle  of 
pronotum  noncarinate;  basal  fovea  small,  shal- 
low; side  margins  shallowly  sinuate.  Interneurs 
effaced  apicolaterally  and  laterally,  striatopunc- 


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TABLE  13.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  1 1.  Bembidion  aeger  n.sp.;  map  Figure  69. 


Lat./Long. 


Elev.  Orig.  Month  No. 

(m)  deposit.  coll.  spec. 


09°35'N 

83°45'W 

3335 

GRNO 

Jan 

10 

09°33'N 

83°45'W 

3416 

USNM 

Mar 

24 

09°32'N 
83°46'W 

3141 

USNM 

Mar 

1 

09°32'N 
83°46'W 

3203 

USNM 

Mar 

6 

09°32'N 
83°46'W 

3355 

USNM 

Jun 

2 

09°32'N 
83°46'W 

1 

UAIC 

Mar 

1 

Total  specimens  examined: 

44 

tulate,  punctulae  small  and  sparse.  Dorsal  mi- 
crosculpture  of  shallowly  engraved  transverse 
meshes,  nearly  effaced  from  elytra  (Fig.  47),  ef- 
faced from  head  and  pronotum.  Male  median 
lobe  as  in  Figure  32.  Size:  see  Tables  2,  3,  and 
4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  69; 
known  only  from  the  upper  slopes  of  Cerro  de 
la  Muerte  and  adjacent  ridges. 

NATURAL  HISTORY  NOTES. — These  beetles 
live  microsympatrically  with  B.  vulcanium,  B. 
lavernae,  and  B.  edwardsi,  on  Cerro  de  la 
Muerte  and  its  adjacent  ridges  at  high  elevations 
in  the  Subparamo  plant  association  (Janzen 
1973),  Montane  Rain  Forest  of  Holdridge  (1971), 
between  3200  m  and  3400  m.  They  were  col- 
lected in  January,  March,  and  June;  teneral 
specimens  were  not  found.  All  individuals  stud- 
ied have  small  nonveined  pads  in  place  of  flight 
wings. 

TYPE-MATERIAL. — Holotype  3 ,  COSTA  RICA,  Cartago,  Cer- 
ro de  la  Muerte,  09°32'N,  083°46'W,  June  (Erwin  &  Erwin) 
(UASM),  ADP  017224.  Paratypes:  256,  189,  same  data  as 
holotype  or  as  listed  in  Table  13  and  deposited  as  USNM-22, 
UASM-2,  CAS-2,  MCZ-2,  AMNH-2,  BMNH-2,  MNHP-2, 
JNEG-2,  UAIC-1,  GRNO-6. 

DERIVATION  OF  TAXON  NAME. — Latin,  ae- 
ger, meaning  sick  or  troubled,  in  reference  to 
the  small  slight  form  of  these  beetles. 

12.  Bembidion  chiriqui,  new  species 

(Figures  14,  33,  48,  69) 

DIAGNOSTIC  COMBINATION. — Eyes  promi- 
nent, produced  well  beyond  outline  of  head  cap- 


sule; form  short  and  subconvex;  elytron  with  all 
interneurs  present,  though  finer  laterally,  all  eas- 
ily traceable  beyond  disc,  broadly  ovoid;  prono- 
tum broad,  with  abruptly  sinuate  side  margins, 
sides  straight  for  only  a  short  distance,  hind  an- 
gle acute;  metasternum  with  intercoxal  process 
broadly  margined,  bead  widely  removed  from 
apex.  Color  and  luster:  dorsum  piceous,  nearly 
black;  scape  and  legs  testaceous,  otherwise  ap- 
pendages infuscated;  venter  piceous;  surfaces 
shiny.  Form:  moderately  small  beetles  with 
moderately  broad  head  and  pronotum;  elytra 
with  arcuate  sides;  pronotum  (Fig.  14)  slightly 
wider  than  head  and  about  equal  to  width  of 
elytron,  markedly  convex  and  cordiform,  base 
prolonged  apically  with  sides  straight  before 
acute  hind  angle;  eyes  large  and  prominent. 
Structure:  frontal  furrows  moderately  well  im- 
pressed, slightly  convergent  on  clypeus.  Hind 
angle  of  pronotum  moderately  carinate,  carina 
not  extended  to  base;  basal  fovea  deep,  extend- 
ed to  side  margin.  Interneurs  entire,  impressed 
laterally,  striatopunctulate,  punctulae  small, 
shallowly  impressed,  widely  spaced.  Dorsal  mi- 
crosculpture  of  elytron  (Fig.  48)  of  moderately 
impressed,  dense,  transverse  lines,  on  head  of 
shallowly  engraved  slightly  transverse  meshes, 
effaced  from  pronotum.  Male  median  lobe  as  in 
Figure  33.  Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  69; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — These  beetles 
were  found  commonly  at  2220  m  altitude  in 
Lower  Montane  Wet  Forest  where  there  was 
sparse  ground  vegetation.  Specimens  were  col- 
lected in  June;  teneral  specimens  were  not 
found.  These  beetles  are  found  independent  of 
water  in  oak  forest  on  finely  textured  soil  under 
leaf  litter.  All  individuals  studied  have  small 
nonveined  pads  in  place  of  flight  wings. 

TYPE-MATERIAL. — Holotype  6 ,  PANAMA,  Chiriqui,  8.0  km 
w  Boquete,  near  Quebrada  Emporio,  08°47'N,  082°30'W,  June 
(Erwin  &  Erwin)  (USNM),  ADP  017618.  Paratypes:  33d ,  109  , 
same  data  as  holotype  and  deposited  as  USNM-29,  UASM-2, 
CAS-2,  MCZ-2,  AMNH-2,  BMNH-2,  MNHP-2,  JNEG-2. 

DERIVATION  OF  TAXON  NAME. — Spanish, 
Chiriqui,  after  the  Volcan  de  Chiriqui  on  which 
these  beetles  live. 

The  nahuala  group 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  without  border;  elytral  interval  3  non- 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


477 


foveate;  male  median  lobe  without  ostium  flags; 
dorsal  microsculpture  absent. 

These  beetles  are  independent  of  running  or 
standing  water  and  occur  in  Subparamo  grass- 
lands of  volcanic  highlands.  The  single  species 
included  in  this  group  and  the  one  in  thefraniae 
group  are  externally  quite  dissimilar,  although 
both  are  similar  to  diverse  members  of  the  sub- 
genus  Peryphus  or  vernale  group  discussed 
above.  However,  males  of  both  nahuala  and 
franiae  have  very  similar  genitalia.  Further 
study  of  these  forms  and  other  related  ones  from 
outside  the  geographic  area  of  this  coverage  will 
need  to  be  undertaken  to  resolve  their  relation- 
ships. 

13.  Bembidion  nahuala,  new  species 

(Figures  15,  34,  49,  70) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  unmargined;  dorsal  microsculpture  ef- 
faced; pronotum  small.  Color  and  luster:  dor- 
sum  piceous;  tibiae,  tarsomeres,  and  antenno- 
meres  1  to  3  testaceous,  otherwise  appendages 
infuscated;  venter  piceous;  surfaces  shiny. 
Form:  small  beetles  with  narrow,  small  head 
and  pronotum,  and  broad  elytra;  pronotum  (Fig. 
15)  about  equal  to  head  in  width,  both  slightly 
narrower  than  elytron;  elytron  either  long  with 
squared  humerus  and  slightly  arcuate  sides,  or 
short  with  rounded  humerus  and  markedly  ar- 
cuate sides;  eyes  large  and  prominent.  Struc- 
ture: frontal  furrows  well  impressed,  rugose, 
parallel,  prolonged  on  clypeus.  Hind  angle  of 
pronotum  acute,  markedly  carinate;  basal  fovea 
large,  deep,  extended  to  basal  margin;  lateral 
margin  evenly  sinuate.  Interneurs  nearly  effaced 
laterally,  represented  by  minute  punctulae,  ef- 
faced apically,  discally  finely  striatopunctulate. 
Dorsal  microsculpture  absent  (Fig.  49).  Male 
median  lobe  as  in  Figure  34.  Size:  see  Tables  2, 
3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  70; 
known  only  from  the  Guatemalan  highlands. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  independent  of  running  or  standing  water 
and  occur  in  Subparamo  grasslands  of  volcanic 
highlands  in  Lower  Montane  Wet  Forest  be- 
tween 2100  m  and  3100  m.  Specimens  were  col- 
lected in  May,  June,  and  August;  teneral  spec- 
imens were  not  found.  At  the  type-locality  these 
beetles  were  found  among  bunch  grass  where 
there  was  little  remaining  forest  due  to  activities 
of  man.  Specimens  were  collected  from  beneath 


TABLE  14.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  13.  Bembidion  nahuala  n.sp.;  map  Figure  70. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month 
coll. 

No. 
spec. 

15°12'N 
90°18'W 

3052 

USNM 

May 

3 

15°04'N 
91°52'W 

2800-2900 

USNM 

May 

50 

14°58'N 
91°46'W 

2200 

USNM 

May 

6 

14°46'N 
91°28'W 

2170 

USNM 

Aug 

1 

14°20'N 
90°31'W 

2105 

USNM 

Jun 

1 

Total  specimens  examined: 

61 

stones  along  the  roadway.  The  species  is  di- 
morphic with  respect  to  wing  length;  brachyp- 
terous  individuals  have  only  a  small  veinless  pad 
in  place  of  the  flight  wing  and  constitute  a  large 
percentage  of  the  populations  sampled  (Table  1). 

TYPE-MATERIAL. — Holotype  cJ ,  GUATEMALA,  San  Marcos, 
27.0  km  NW  San  Marcos,  15°04'N,  091°52'W,  May  (Erwin  & 
Erwin)  (USNM),  ADP  013024.  Paratypes:  18cJ,  429,  same 
data  as  holotype  or  as  listed  in  Table  14  and  deposited  as 
USNM-40,  UASM-2,  CAS-4,  MCZ-4,  AMNH-2,  BMNH-2, 
MNHP-2,  JNEG-2,  FMNH-2. 

DERIVATION  OF  TAXON  NAME. — Uto-aztec- 
an,  Nahuala,  after  the  Rio  Nahuala  where  the 
type-series  was  collected;  the  word  derives  from 
the  general  name  for  the  indigenous  peoples  of 
the  region,  the  Nahuatl,  of  which  the  Aztecs 
were  part. 

Thefraniae  group 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  without  border;  elytral  interval  3  non- 
foveate;  male  median  lobe  with  ostium  flags; 
dorsal  microsculpture  present. 

These  beetles  are  independent  of  running  or 
standing  water  and  occur  in  litter  in  Lower  Mon- 
tane Moist  Forest.  The  single  species  included 
in  this  group  and  the  one  in  the  nahuala  group 
are  externally  quite  dissimilar,  although  both  are 
similar  to  diverse  members  of  the  subgenus 
Peryphus  or  vernale  group  discussed  above. 
However,  males  of  both  nahuala  and  franiae 
have  very  similar  genitalia.  Further  study  of 
these  forms  and  other  related  ones  from  outside 
the  geographic  area  of  this  coverage  will  need 
to  be  undertaken  to  resolve  their  relationships. 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


FIGURES  40  to  53.  Microsculpture  of  elytron,  left,  dorsal  aspect  posterior  to  Ed3,  3rd  interval,  ca.  lOOOx,  males.  40.  B. 
rogersi  Bates,  7.5  km  south  of  Ojo  de  Agua,  Cartago,  Costa  Rica.  41.  B.  mexicanum  Dejean,  22.0  km  west  of  San  Pedro 
Sacatepequez  at  Rio  Nahuala.  42.  B.  vernale  Bates,  San  Isidro  de  Coronado,  San  Jose,  Costa  Rica.  43.  B.  satellites  Bates, 
Chomogo  area,  10°18'N,  084°47'W,  Alajuela,  Costa  Rica.  44.  B.  quetzal  n.sp.,  25.0  km  south  of  Totonicapan,  Huehuetenango, 
Guatemala.  45.  B.  diabola  n.sp.,  Volcan  Irazii,  Cartago,  Costa  Rica.  46.  B.  vulcanium  Darlington,  7.5  km  south  of  Ojo  de 
Agua,  Cartago,  Costa  Rica.  47.  B.  aeger  n.sp.,  Cerro  Buenavista,  San  Jose,  Costa  Rica.  48.  B.  chiriqui  n.sp.,  8.0  km  west  of 
Boquete,  Chiriqui,  Panama.  49.  B.  nahuala  n.sp.,  27.0  km  northwest  of  San  Marcos,  San  Marcos,  Guatemala.  50.  B.  aratum 
LeConte,  La  Lima,  Cortes,  Honduras.  51.  B.  cartes  n.sp..  La  Lima,  Cortes,  Honduras.  52.  B.  sparsum  Bates,  Tikal,  Del 
Peten,  Guatemala.  53.  B.  ixtatan  n.sp.,  27.0  miles  [43.5  km]  west  of  Solola,  Solola,  Guatemala. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


479 


14.  Bembidion  franiae,  new  species 

(Figures  16,  35,  70) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  between  mesocoxae  entirely  unmar- 
gined;  elytron  bipunctate,  punctures  around 
dorsal  setae  deep  and  small,  not  extended  be- 
yond middle  of  adjacent  interval;  dorsal  surface 
with  well-developed  microsculpture,  that  of  ely- 
tra of  finely  impressed  lines  or  wide  meshes,  sur- 
face subiridescent,  on  head  clearly  and  evenly 
isodiametric,  slightly  granular,  more  evenly 
transverse  and  less  impressed  on  pronotum. 
Color  and  luster:  dorsum  piceous;  outer  anten- 
nomeres  and  penultimate  palpomeres  infuscat- 
ed,  otherwise  appendages  testaceous;  venter  pi- 
ceous; surfaces  shiny,  elytra  subiridescent. 
Form:  moderately  small  beetles  for  genus,  with 
large  head  and  pronotum  in  proportion  to  elytra; 
pronotum  (Fig.  16)  wider  than  head  and  elytron; 
eyes  large  and  prominent.  Structure:  frontal  fur- 
rows shallow  and  broad,  prolonged  on  clypeus; 
anterior  supraorbital  pore  deeply  foveate.  Hind 
angle  of  pronotum  markedly  carinate;  basal  fo- 
vea  large  and  deep;  lateral  margins  sinuate, 
straight  for  some  distance  behind  sinuation.  In- 
terneurs  of  large  punctulae  laterally,  effaced  at 
extreme  apex;  disc  markedly  striatopunctulate, 
punctulae  wide-spaced,  not  coarsely  impressed. 
Dorsal  microsculpture  of  elytra  finely  engraved, 
markedly  transverse  meshes,  of  head  isodia- 
metric, slightly  granular  meshes,  and  on  prono- 
tum well-engraved  transverse  meshes.  Size:  see 
Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  70; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  independent  of  running  or  standing  water 
and  occur  in  oak  forests  in  volcanic  highlands 
in  Lower  Montane  Wet  Forest  at  2500  m  and 
2600  m.  Specimens  were  collected  in  August; 
the  paratype  is  teneral.  At  the  type-locality  these 
beetles  were  found  in  oak  leaf  litter.  The  species 
is  probably  flightless,  judging  from  the  markedly 
developed  state  of  brachyptery  in  the  two 
known  individuals  which  have  only  a  small  vein- 
less  pad  in  place  of  the  flight  wing. 

TYPE-MATERIAL. — Holotype  c?,  GUATEMALA,  Huehuete- 
nango,  4.8  km  E  San  Mateo  Ixtatan,  15°50'N,  091°27'W,  Aug. 
(Ball,  Frania,  &  Whitehead)  (USNM),  ADP  026917.  Paratype: 
1 2 ,  same  data  as  type  (UASM). 

DERIVATION  OF  TAXON  NAME. — I  take  plea- 
sure in  naming  this  species  in  honor  of  one  of 
the  collectors  of  the  types,  Henry  Frania. 


The  incrematum  group 

(Subgenus  Eupetedromus  Netolitzky,  1911:190.  Type-species 
Carabus  dentellus  Thunberg,  1787:50,  by  subsequent  desig- 
nation of  Lindroth  1963:348.) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  not  bordered;  elytra  iridescent  from 
densely  transverse  microsculpture  or  micro- 
sculpture  absent;  male  genitalia  similar  to  those 
in  members  of  subgenus  Nothaphus  (Fig.  36; 
see  also  Lindroth  1963:367). 

These  beetles  are  hygrophilous,  occurring 
along  streams  or  standing  water  in  fine  gravel 
and  sand,  or  on  silt.  Previously,  this  group  was 
regarded  as  Holarctic;  the  addition  to  the  group 
of  B.  aratum  of  the  southwestern  United  States, 
Mexico,  and  Central  America  extends  the 
group's  range  considerably.  All  species  are  fully 
winged  and  occur  in  the  lowland  or  adjacent  low 
hills.  The  male  median  lobe  of  the  species  cov- 
ered herein  has  the  pigmented  ostium  flag  char- 
acteristic of  North  and  Central  American  Bem- 
bidion (Notaphus)  species,  and  the  endophallus 
strongly  suggests  relationship  with  this  group. 

15.  Bembidion  aratum  LeConte 

(Figures  17,  36,  50,  71) 

Bembidium  aratus  LECONTE,  1852:189.  [Lectotype  2,  ARI- 
ZONA, Gila  River  Valley  (MCZ),  designated  by  Erwin 
1982.] 

Bembidium  scintillans  BATES,  1882:150.  [Lectotype  9,  MEX- 
ICO, Capulapam,  17°18'N,  096°27'W  (BMNH),  designated 
by  Erwin  1982.] 

Bembidion  vinnulum  CASEY,  1918:116.  [Lectotype  9,  ARI- 
ZONA (Southern)  (USNM),  designated  by  Erwin  1982.] 

Bembidion  definitum  CASEY,  1918:166.  [Holotype  <J ,  ARIZO- 
NA, Tucson,  32°13'N,  110°57'W  (USNM).] 

DIAGNOSTIC  COMBINATION. — Elytron  with 
setae  Ed3  and  Ed5  in  interval  3  and  not  touching 
adjacent  interneurs;  elytron  without  microsculp- 
ture, markedly  shiny.  Color  and  luster:  dorsum 
piceous;  elytra  maculate,  tibial  base,  femur 
apex,  distal  antennomeres,  penultimate  palpo- 
meres infuscated,  otherwise  appendages  testa- 
ceous; venter  piceous;  surfaces  markedly  shiny, 
elytra  with  faint  metallic  cast.  Form:  medium- 
sized  beetles  for  genus  with  narrow  head  and 
pronotum  and  long-tapered  elytra;  pronotum 
(Fig.  17)  slightly  wider  than  head,  markedly  con- 
vex, about  equal  to  width  of  elytron;  eyes  large 
and  prominent.  Structure:  frontal  furrows  shal- 
low, broad,  parallel,  extended  to  clypeus.  Hind 
angle  of  pronotum  markedly  carinate;  basal  fo- 
vea  small,  deep,  extended  to  basal  margin;  lat- 
eral margins  shallowly  sinuate.  Interneurs  en- 


480 


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TABLE  15.    LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  15.  Bembidion  aratum  LeConte;  map  Figure  71. 


TABLE  16.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  16.  Bembidion  cortes  n.sp.;  map  Figure  71. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month 
coll. 

No. 
spec. 

15°30'N 
86°35'W 

0140-0160 

UAIC 

Jun 

1 

15°26'N 
87°55'W 

0020 

FDAG 

Jun 

2 

15°26'N 
87°55'W 

0020 

UASM 

Jun 

1 

15°25'N 
91°43'W 

1440 

USNM 

Aug 

2 

Total  specimens  examined: 

6 

tire,  deeply  impressed,  striatopunctulate 
anteriorly,  striate  posteriorly.  Dorsal  micro- 
sculpture  of  head  of  shallowly  impressed,  mod- 
erate-sized, slightly  transverse  meshes,  effaced 
from  pronotum  and  elytron  (Fig.  50).  Male  me- 
dian lobe  as  in  Figure  36.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  71; 
southwestern  United  States  to  Honduras  along 
the  central  and  eastern  parts  of  the  continent. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  from  sea  level  to  middle  elevations  in 
Tropical  Dry,  Tropical  Very  Dry,  and  Subtrop- 
ical Dry  forests  up  to  1500  m.  Specimens  were 
collected  in  June  and  August;  no  teneral  speci- 
mens were  found.  These  beetles  occur  at  the 
edge  of  moderately  large  streams  in  gravel  and 
under  stones  where  there  is  some  silt  and  at  least 
some  sparse  vegetation.  All  known  specimens 
are  fully  winged;  they  were  attracted  to  UV  light 
traps  in  Mexico,  Honduras,  and  Guatemala,  in- 
dicating that  the  beetles  fly. 

MATERIAL  EXAMINED. — Types  (see  above)  and  6  speci- 
mens (Table  15). 

The  dorsale  group 

(Subgenus  Notaphus  Stephens,  1828:2.  Type-species  Carabus 
varius  Olivier,  1795:110,  by  subsequent  designation  of  West- 
wood  1840:7.) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered;  elytra  and  fore- 
body  with  isodiametric  microsculpture;  elytra 
with  light-colored  mosaic  pattern  on  dark  back- 
ground; male  genitalia  uniform  and  similar  to 
those  in  members  of  subgenus  Eupetedromus 
(Fig.  36,  see  also  Lindroth  1963:367). 

These   beetles   are   hygrophilous,    occurring 


Lat./Long. 

Elev. 

(m) 

Orig. 
deposit. 

Month         No. 
coll.         spec. 

15°26'N 
87°55'W 

0020 

FDAG 

Jun              18 

15°26'N 
87°55'W 

0020 

UASM 

Jun             14 

Total  specimens  examined:     32 

along  streams  or  standing  water  in  fine  gravel 
and  sand  or  among  stones,  or  on  sea  beaches. 
Previously,  this  group  was  regarded  as  predom- 
inantly Holarctic,  however,  several  species  oc- 
cur in  the  Tropics  and  in  South  Temperate  re- 
gions, mainly  on  beaches.  All  species  are  fully 
winged  and  occur  in  the  lowland  or  adjacent  low 
hills.  The  male  median  lobes  of  the  species  cov- 
ered herein  have  the  pigmented  ostium  flag  char- 
acteristic of  North  American  Bembidion  (No- 
taphus)  species. 

16.  Bembidion  cortes,  new  species 

(Figures  18,37,  51,71) 

DIAGNOSTIC  COMBINATION. — Elytron  with 
setae  Ed3  and  Ed5  in  interval  3,  not  touching 
adjacent  interneurs,  and  with  perfectly  isodia- 
metric  and  well-engraved  microsculpture; 
pronotum  broad,  moderately  convex,  much  wid- 
er than  head  across  eyes,  side  margins  not  mark- 
edly arcuate  nor  much  convergent  behind,  shal- 
lowly  sinuate  before  hind  angle.  Color  and 
luster:  dorsum  of  elytron  with  mosaic  color  pat- 
tern, head  and  pronotum  dark  green;  distal  an- 
tennomeres  infuscated,  otherwise  appendages 
flavous  or  testaceous;  venter  rufopiceous  or  pi- 
ceous,  in  some  specimens  apex  of  abdomen  pal- 
er; surface  of  elytra  shiny,  forebody  metallic. 
Form:  medium-sized  beetles  for  this  genus  with 
head,  pronotum,  and  elytra  broad;  pronotum 
(Fig.  18)  somewhat  wider  than  head  and  elytron; 
eyes  large  and  prominent.  Structure:  frontal  fur- 
rows very  shallow  and  broad,  extended  on  clyp- 
eus.  Hind  angle  of  pronotum  markedly  carinate; 
basal  fovea  broad  and  deep;  lateral  margins  shal- 
lowly  sinuate.  Interneurs  entire,  deeply  im- 
pressed throughout  their  length,  striatopunctu- 
late  anteriorly,  striate  posteriorly.  Dorsal 
microsculpture  of  elytron  nearly  isodiametric, 
with  some  transverse  meshes  (Fig.  51),  that  of 
head  and  pronotum  perfectly  isodiametric, 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


481 


slightly  granulate,  deeply  engraved.  Male  me- 
dian lobe  as  in  Figure  37.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  71; 
southern  Mexico  north  at  least  to  Vera  Cruz, 
south  to  Honduras. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  low  elevations  in  Tropical  Dry  For- 
ests at  20  m.  Specimens  were  collected  in  June 
and  December;  no  teneral  specimens  were 
found.  All  known  specimens  are  fully  winged; 
they  were  attracted  to  UV  light  traps  in  Mexico 
and  Honduras,  indicating  that  the  beetles  fly. 

TYPE-MATERIAL. — Holotype  <J,  HONDURAS,  Cortes,  La 
Lima,  15°26'N,  087°55'W,  June  (Blanton,  Brace,  &  Woodruff) 
(FDAG),  ADP  047025.  Paratypes:  96 ,  22$  ,  same  data  as  ho- 
lotype  or  as  listed  in  Table  16  and  deposited  as  USNM-12, 
UASM-2,  CAS-2,  MCZ-2,  BMNH-2,  MNHP-2,  JNEG-2, 
FDAG-7. 

DERIVATION  OF  TAXON  NAME. — Spanish, 
Cortes,  after  the  sea-faring  explorer  from  Spain 
who  touched  the  Middle  American  coast  and 
Caribbean  islands  in  so  many  places;  in  refer- 
ence to  the  probable  habitat  and  distribution  of 
these  beetles. 

17.  Ik-iii  I)  id  ion  spar  sum  Bates 

(Figures  19,  38,  52,  71) 

Bembidium  sparsum  BATES,  1882:151.  [Lectotype  d,  MEXI- 
CO, Oaxaca  (BMNH),  designated  by  Erwin  1982.] 

DIAGNOSTIC  COMBINATION. — Elytron  with 
setae  Ed3  and  Ed5  in  interval  3  and  not  touching 
adjacent  interneurs,  and  with  perfectly  isodia- 
metric  and  well-engraved  microsculpture; 
pronotum  narrow,  about  as  wide  as  head  across 
eyes,  markedly  constricted  behind.  Color  and 
luster:  dorsum  of  elytron  with  mosaic  color  pat- 
tern, head  and  pronotum  dark  green;  distal  an- 
tennomeres  infuscated,  otherwise  appendages 
flavous  or  testaceous;  venter  rufopiceous  or  pi- 
ceous,  in  some  specimens  apex  of  abdomen  pal- 
er; surface  of  elytra  shiny,  forebody  metallic. 
Form:  medium-sized  beetles  for  this  genus  with 
head,  pronotum,  and  elytra  broad;  pronotum 
(Fig.  19)  narrow,  equal  in  width  to  head  and 
elytron;  eyes  large  and  prominent.  Structure: 
frontal  furrows  very  shallow  and  broad,  extend- 
ed on  clypeus.  Hind  angle  of  pronotum  mark- 
edly carinate;  basal  fovea  broad  and  deep;  lat- 
eral margins  deeply  sinuate.  Interneurs  entire, 
deeply  impressed  throughout  their  length,  stria- 
topunctulate  anteriorly,  striate  posteriorly.  Dor- 


sal microsculpture  of  elytron  nearly  isodiamet- 
ric,  with  some  transverse  meshes  (Fig.  52),  that 
of  head  and  pronotum  perfectly  isodiametric, 
slightly  granulate,  deeply  engraved.  Male  me- 
dian lobe  as  in  Figure  38.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  71; 
most  of  Middle  America  from  Sonora,  Mexico, 
south  to  at  least  Nicaragua  and  along  the  east 
coast  at  Vera  Cruz,  Mexico,  as  well  as  on  the 
islands  of  Jamaica  and  Hispaniola  (Haiti),  and 
in  Colombia.  Curiously,  there  are  no  records 
from  Costa  Rica  or  Panama. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  low  elevations  in  Tropical  Dry, 
Tropical  Very  Dry,  and  Tropical  Moist  forests 
between  20  m  and  500  m.  Specimens  were  col- 
lected in  June,  July,  August,  and  September  on 
the  mainland,  February  on  Jamaica,  and  Sep- 
tember on  Hispaniola;  some  teneral  specimens 
were  found  on  Jamaica  and  Hispaniola,  and 
some  of  the  September  mainland  specimens 
were  also  teneral.  These  beetles  frequent  low- 
lying  areas  on  dry  sand  which  are  adjacent  to  in- 
termittently wet  streams;  the  general  vegetation 
is  thorn  scrub.  All  known  specimens  are  fully 
winged;  they  were  attracted  to  UV  light  traps  in 
many  localities,  indicating  that  the  beetles  fly. 
This  together  with  their  occurrence  in  the  Great- 
er Antilles  and  their  preference  for  intermittent- 
ly wet  areas  indicate  they  are  highly  vagile  dis- 
persants. 

MATERIAL  EXAMINED. — Lectotype  (see  above)  and  17 
specimens  (Table  17). 

18.  Bembidion  armuelles,  new  species 

(Figures  20,  71) 

DIAGNOSTIC  COMBINATION. — Elytron  with 
setae  Ed3  and  Ed5  in  interval  3  and  not  touching 
adjacent  interneurs,  and  with  perfectly  isodia- 
metric and  well-engraved  microsculpture; 
pronotum  very  broad  and  convex  with  abruptly 
sinuate  lateral  margins  basally,  much  wider  than 
head  across  eyes,  markedly  constricted  behind. 
Color  and  luster:  dorsum  of  elytron  with  mosaic 
color  pattern,  head  and  pronotum  dark  green; 
distal  antennomeres  infuscated,  otherwise  ap- 
pendages flavous  or  testaceous;  venter  rufopi- 
ceous or  piceous,  in  some  specimens  apex  of 
abdomen  paler;  surfaces  of  elyta  shiny,  fore- 
body  metallic.  Form:  medium-sized  beetles  for 
this  genus  with  head,  pronotum,  and  elytra 


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TABLE  17.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  17.  Bembidion  sparsum  Bates;  map  Figure  71. 


Lat./Long. 


Elev.  Orig.          Month        No. 

(m)  deposit.          coll.         spec. 


17°25'N 
88°29'W 

0000-0300 

USNM 

Jun 

1 

17°20'N 
89°39'W 

0250 

USNM 

Jun 

3 

17°20'N 
88°33'W 

0000-0300 

USNM 

Jun 

1 

17°15'N 
88°45'W 

0000-0300 

USNM 

Jun 

1 

15°44'N 
87°27'W 

0000-0020 

UMAA 

Mar 

2 

15°26'N 
87°55'W 

0020 

UASM 

Jun 

4 

13°52'N 
90°05'W 

0030-0040 

USNM 

Jul 

2 

13°36'N 
89°50'W 

0000-0010 

UASM 

May 

1 

12°36'N 
86°02'W 

0457 

USNM 

Jul 

1 

12°23'N 
86°03'W 

0122 

USNM 

Jun 

1 

Total  specimens  examined: 

17 

broad;  pronotum  (Fig.  20)  very  broad  and  con- 
vex, wider  than  head  and  elytron;  eyes  large  and 
prominent.  Structure:  frontal  furrows  very  shal- 
low and  broad,  extended  on  clypeus.  Hind  angle 
of  pronotum  markedly  carinate;  basal  fovea 
broad  and  deep;  lateral  margins  deeply  sinuate. 
Interneurs  entire,  deeply  impressed  throughout 
their  length,  striatopunctulate  anteriorly,  striate 
posteriorly;  intervals  convex.  Dorsal  micro- 
sculpture  of  elytron  nearly  isodiametric,  with 
some  transverse  meshes,  that  of  head  and 
pronotum  perfectly  isodiametric,  slightly  gran- 
ulate, deeply  engraved.  Male  unknown.  Size: 
see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  71; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — The  town  of  Ar- 
muelles  is  a  port  on  the  Pacific  Ocean  with  ex- 
tensive sea  beach;  behind  the  town  lies  Tropical 
Moist  Forest.  I  do  not  know  the  habitat  in  which 
the  unique  specimen  was  collected;  however,  it 
is  fully  winged,  it  was  collected  in  June,  and  it 
is  not  teneral. 

TYPE-MATERIAL. — Holotype  9 ,  PANAMA,  Chiriqui,  Puerto 
Armuelles,  08°17'N,  082°52'W,  June  (Bierig)  (MCZ),  ADP 
003 142. 


DERIVATION  OF  TAXON  NAME. — Spanish, 
Armuelles,  the  name  of  the  town  near  which  the 
holotype  was  collected. 

The  a/fine  group 

(Subgenus  Furcacampa  Netolitzky,   1931:158.  Type-species 
Bembidium  affine  Say,  1823:86,  by  original  designation.) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered,  margin  removed 
from  apex;  elytra  and  forebody  with  isodiamet- 
ric microsculpture;  elytra  with  light-colored  mo- 
saic pattern  on  dark  background;  male  genitalia 
similar  to  those  in  members  of  versicolor  group 
(fig.  191  in  Lindroth  1963:379). 

These  beetles  are  hygrophilous,  occurring 
along  streams  or  standing  water  on  fine  clay  or 
sand,  at  the  margins  of  marshy  pools,  or  on  lake 
beaches.  Previously,  this  group  was  regarded  as 
Nearctic,  however,  the  new  species  described 
below  from  Panama  extends  the  range  consid- 
erably. Both  species  of  the  group  are  fully 
winged  and  occur  in  the  lowland  or  adjacent  low 
hills. 

19.  Bembidion  barrensis,  new  species 

(Figures  22b,  39b,  74) 

DIAGNOSTIC  COMBINATION. — Frontal  fur- 
rows shallow,  single,  and  nearly  parallel  be- 
tween eyes,  convergent  and  doubled  on  clypeus. 
Color  and  luster:  dorsum  of  elytron  with  mosaic 
color  pattern,  head  and  pronotum  dark  piceous; 
appendages  testaceous  (on  holotype,  outer  an- 
tennomeres  missing,  palpomeres  missing);  ven- 
ter piceous;  surfaces  of  elytra  shiny,  forebody 
slightly  metallic.  Form:  small-sized  for  genus 
with  narrow  forebody  and  elytra;  elytra  with 
square  humeri;  pronotum  (Fig.  lib)  narrower 
than  head,  about  equal  in  width  to  elytron;  eyes 
large  and  prominent.  Structure:  frontal  furrows 
well  impressed,  slightly  rugose,  almost  parallel 
between  eyes,  angularly  prolonged  and  doubled 
on  clypeus.  Hind  angle  of  pronotum  acute,  with 
feebly  developed,  short,  sharp  carinae;  basal 
fovea  large  and  deep;  side  margins  abruptly  and 
markedly  sinuate.  Interneurs  entire,  though 
much  shallower  or  effaced  apicolate rally,  stria- 
topunctulate, punctulae  medium-sized,  coarse, 
and  close-spaced.  Dorsal  microsculpture  of 
large  transverse  meshes  on  apex  of  elytron,  head 
and  pronotum,  effaced  from  disc  of  elytron. 
Male  median  lobe  as  in  Figure  3>9b.  Size:  see 
Tables  2,  3,  and  4. 


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483 


GEOGRAPHICAL  DISTRIBUTION. — Figure  74; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — The  single  spec- 
imen was  found  in  Tropical  Moist  Forests  at  28 
m.  It  was  collected  in  October  and  is  not  teneral. 
It  is  fully  winged.  The  habitat  consists  of  marshy 
lake  shore  (on  the  back  side  of  the  island  op- 
posite the  Canal)  with  rich  growth  of  Calathea 
and  other  emergent  broadleaf  plants  growing  in 
shallow,  but  rich,  organic  soil  on  top  of  red  la- 
terite.  The  marsh  is  heavily  shaded  in  this  area. 
The  beetle  was  in  a  thin  layer  of  leaves  less  than 
a  meter  from  the  water. 

TYPE-MATERIAL. — Holotype  6 ,  PANAMA,  Canal  Zone,  Bar- 
ro  Colorado  Island,  09°10'N,  079°51'W,  Oct.  (Erwin  &  Erwin) 
(USNM),  ADP041711. 

DERIVATION  OF  TAXON  NAME. — Spanish, 
barro,  meaning  clay  or  mud  and  part  of  the 
name  of  the  place  where  the  holotype  was  col- 
lected. 

The  championi  group 

(Subgenus  Cyclolopha  Casey,  1918:144.  Type-species  Bem- 
bidium  sphaeroderum  Bates,  1882:147,  here  designated.) 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered,  margin  not  re- 
moved from  apex;  pronotum  markedly  lobed 
basally,  hind  angle  absent  or  markedly  obtuse; 
frontal  furrows  deep,  markedly  convergent,  pro- 
longed on  clypeus. 

Little  is  known  regarding  the  life  history  and 
habits  of  these  beetles.  Few  specimens  are  avail- 
able for  study.  I  collected  one  individual  of  the 
group  in  Mexico  from  beneath  a  rosette  of  pros- 
trate grasses  on  a  volcanic  hillside  in  the  Trans- 
verse Volcanic  Belt.  There  was  no  water  close 
by.  A  partially  complete  manuscript  by  G.  G. 
Perrault  (in  litt.)  indicates  there  are  eight  species 
in  the  group,  distributed  from  Colorado,  USA, 
to  Guatemala. 

20.  Bembidion  cyclodes  Bates 

(Figure  72) 

Bembidium  cyclodes  BATES,  1884:290.  [Lectotype  9,  MEXI- 
CO, Oaxaca  (BMNH),  selected  and  labelled  by  Perrault, 
herewith  designated.] 

Bembidium  aztecanum  CASEY,  1918:145.  [Lectotype  cJ,  MEX- 
ICO, Cuernavaca  (USNM),  selected  and  labelled  by  Per- 
rault, here  designated.] 

DIAGNOSTIC  COMBINATION. — Pronotum  with 
hind  angle  rounded  with  small  angle  along  mar- 


TABLE  18.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  20.  Bembidion  cyclodes  Bates;  map  Figure  72. 


Lat./Long. 

Elev. 
(m) 

Orig. 
deposit. 

Month          No. 
coll.           spec. 

[Types 
from 
Mexico] 

1 

BMNH, 

USNM 

?                     2 

15°23'N 
91°02'W 

1829 

AMNH 

Aug                 1 

Total  specimens  examined:      3 

gin  at  posterior  seta,  base  markedly  lobed;  ely- 
tron with  interneur  7  absent,  interneur  6  extend- 
ed to  at  least  middle.  Color  and  luster:  dorsum 
of  elytra  and  propleura  rufous;  distal  antenno- 
meres  infuscated,  otherwise  appendages  testa- 
ceous; head,  pronotum,  and  venter  piceous;  sur- 
faces markedly  shiny.  Form:  small  beetles  with 
broad  forebody  and  elytra;  pronotum  slightly 
wider  than  head  or  elytron;  eyes  markedly  large 
and  prominent.  Structure:  frontal  furrows  sul- 
cate,  narrow,  convergent,  extended  to  clypeal 
setigerous  pore.  Hind  angle  of  pronotum  mark- 
edly obtuse,  nearly  absent;  dorsum  markedly 
convex;  lateral  margins  markedly  arcuate  pos- 
teriorly; base  prominently  lobed.  Interneurs  1  to 
6  each  a  row  of  punctulae  in  basal  half  of  ely- 
tron, 7  absent,  1  striatopunctulate,  in  apical  half 
2  to  6  absent;  punctulae  separated  by  their  own 
diameter.  Dorsal  microsculpture  absent.  Size: 
see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  72; 
known  specifically  from  only  one  locality  in 
Guatemala;  older  records  give  only  "Guat." 
Perrault  recorded  this  species  from  localities  in 
Mexico,  north  as  far  as  Cuernavaca. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  middle  elevations  in  Lower  Mon- 
tane Moist  Forests  between  1450  m  and  1900  m. 
Specimens  were  collected  in  August  and  Octo- 
ber; no  teneral  specimens  were  found.  All 
known  specimens  are  fully  winged. 

MATERIAL  EXAMINED. — Types  (see  above)  and  1  specimen 
(Table  18). 

2 1 .  Bembidion  championi  Bates 

(Figures  2 1,72) 

Bembidium  championi  BATES,  1882:148.  [Lectotype  cJ,  GUA- 
TEMALA, Quiche  Mountains  (MNHP),  selected  and  labelled 
by  Perrault,  here  designated.] 


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TABLE  19.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  21.  Bembidion  championi  Bates;  map  Figure  72. 


TABLE  20.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  22.  Bembidion  ixtatan  n.sp.;  map  Figure  72. 


Elev. 

Orig.          Month 

No. 

Elev. 

Orig. 

Month         No. 

Lat./Long.                   (m) 

deposit.         coll. 

spec. 

Lat./Long. 

(m) 

deposit. 

coll.          spec. 

[Type  from                    9 
Guatemala] 

BMNH            ? 

1 

15°50'N 
91°27'W 

2460 

USNM 

Aug               6 

[Mexico.  Chiapas, 
8.6  mi.  E 

UASM 

2 

14°47'N 
91°15'W 

ca.  2400 

USNM 

Aug                 1 

San  Cristobal] 

Total  specimens  examined:       7 

DIAGNOSTIC  COMBINATION. — Pronotum  with 
hind  angle  rounded,  without  small  angle  along 
margin  at  posterior  seta,  base  markedly  lobed; 
elytron  with  interneurs  6  and  7  present,  short, 
not  extended  beyond  basal  sixth.  Color  and  lus- 
ter: dorsum  of  elytra  and  propleura  rufous;  distal 
antennomeres  infuscated,  otherwise  appendages 
testaceous;  head,  pronotum,  and  venter  pi- 
ceous;  surface  markedly  shiny.  Form:  small 
beetles  with  broad  forebody  and  elytra;  prono- 
tum slightly  wider  than  head  or  elytron;  eyes 
markedly  large  and  prominent.  Structure:  fron- 
tal furrows  sulcate,  narrow,  convergent,  extend- 
ed to  clypeal  setigerous  pore.  Hind  angle  of 
pronotum  markedly  obtuse,  nearly  absent;  dor- 
sum  markedly  convex;  lateral  margins  markedly 
arcuate  posteriorly;  base  prominently  lobed.  In- 
terneurs 1  to  7  each  a  row  of  punctulae  in  basal 
half  of  elytron,  1  striatopunctulate,  in  apical  half 
2  to  7  absent;  punctulae  separated  by  more  than 
their  own  diameter.  Dorsal  microsculpture  ab- 
sent. Size:  see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  72; 
known  specifically  from  only  the  type-locality  in 
Guatemala;  other  records  give  only  "Guat." 
Perrault  recorded  this  species  from  localities  in 
Mexico,  north  as  far  as  Jalisco. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  middle  elevations  in  Lower  Mon- 
tane Moist  Forests  between  2100  m  and  2700  m. 
Specimens  were  collected  in  June,  August,  and 
September;  no  teneral  specimens  were  found. 
All  known  specimens  have  the  wings  reduced  to 
small  pads,  each  of  which  retains  the  bases  of 
two  major  veins. 

MATERIAL  EXAMINED. — Lectotype  (see  above)  and  2  spec- 
imens (Table  19). 


The  cognatum  group 

DIAGNOSTIC  COMBINATION. — Metasternal 
process  completely  bordered,  margin  not  re- 
moved from  apex;  pronotum  not  lobed  basally, 
hind  angles  nearly  square;  frontal  furrows  deep, 
markedly  convergent,  prolonged  on  clypeus; 
microsculpture  effaced  from  pronotum  and  ely- 
tra. 

These  beetles  are  hygrophilous,  occurring 
along  streams  in  fine  gravel  and  sand,  or  in 
mosses.  Previously,  this  group  of  two  species 
was  known  from  only  North  America  and  Mex- 
ico; the  Central  American  one  described  below 
extends  the  group's  range  considerably.  Both 
species  are  fully  winged  and  occur  at  middle  el- 
evations. The  male  median  lobe  of  the  species 
covered  herein  has  two  pigmented  ostium  flags. 

This  group  has  not  been  assigned  to  subgenus. 
Its  members  are  intermediate  between  tran- 
sparens  group  (head  structure)  and  versicolor 
group  (body  structure  and  color  pattern).  The 
unique  male  genitalia  does  not  indicate  relation- 
ship to  either  of  these  mentioned  groups.  Place- 
ment must  await  revisions  of  the  several  species 
groups  with  convergent  frontal  furrows  (see  Er- 
win  and  Kavanaugh  1981). 

22.  Bembidion  ixtatan,  new  species 

(Figures  22a,  39a,  53,  72) 

DIAGNOSTIC  COMBINATION. — Pronotum  with 
hind  angle  squared;  base  not  lobed;  elytron  with 
all  interneurs  coarsely  punctulate  well  beyond 
middle;  frontal  furrows  markedly  convergent; 
microsculpture  effaced  from  elytra  and  prono- 
tum. Color  and  luster:  elytra  piceous,  obscurely 
maculate,  maculae  not  well  defined;  forebody 
and  venter  piceous;  appendages  infuscated;  sur- 
faces markedly  shiny.  Form:  small  beetles  for 
the  genus  with  head  large,  pronotum  narrow, 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


485 


elytra  convex  and  narrow;  pronotum  (Fig.  22a) 
slightly  wider  than  head  and  elytron;  eyes  mod- 
erately large  and  prominent.  Structure:  frontal 
furrows  well  impressed,  markedly  convergent 
toward  clypeus,  doubled  on  clypeus.  Hind  angle 
of  pronotum  nearly  square,  finely  carinate,  ca- 
rinae  markedly  short;  basal  fovea  small  and 
deep;  lateral  margins  shallowly,  abruptly  sin- 
uate. Interneurs  effaced  apicolaterally,  mark- 
edly and  coarsely  punctulate  on  disc.  Dorsal 
microsculpture  effaced  (Fig.  53).  Male  median 
lobe  as  in  Figure  39a.  Size:  see  Tables  2,  3, 
and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  72; 
known  only  from  the  Guatemalan  highlands. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  at  upper  to  middle  elevations  in  Lower 
Montane  Moist  Forests  at  2500  m.  Specimens 
were  collected  in  August;  no  teneral  specimens 
were  found.  These  beetles  live  near  mountain 
streams  where  they  can  be  found  during  the  day 
by  washing  mosses.  All  known  specimens  are 
fully  winged. 

TYPE-MATERIAL. — Holotype  $ ,  GUATEMALA,  Huehuete- 
nango,  4.8  km  E  San  Mateo  Ixtatan,  15°50'N,  091°27'W  (Ball, 
Frania,  &  Whitehead)  (USNM)  ADP  026911.  Paratypes:  2<J, 
49 ,  same  data  as  for  holotype  or  as  listed  in  Table  20  and 
deposited  as  USNM-2,  UASM-2,  CAS-2. 

DERIVATION  OF  TAXON  NAME. — Probably 
Uto-Aztecan,  Ixtatan,  from  the  name  of  the 
town  near  which  the  type-series  was  collected. 

Genus  Geocharidius  Jeannel 

Geocharidius  JEANNEL,  1963a:107.  [Type-species  Anillus  in- 
tegripennis  Bates,  1882:145,  by  original  designation.] 

For  details  on  related  groups  see  Jeannel 
(1937,  1960,  1963a),  and  Taglianti  (1973). 

DIAGNOSTIC  COMBINATION. — Small-sized, 
more  or  less  depressed  or  markedly  convex  bee- 
tles with  narrow  or  broad  elytra,  narrow  fore- 
body,  and  with  medium-length,  slender  append- 
ages. Color  pale  rufotestaceous.  Terminal 
palpomere  subulate;  eyes  absent;  antennomeres 
2  to  11  entirely  pubescent  and  with  numerous 
long  setae,  mentum  with  well-developed  but 
small  tooth;  mandibles  unisetose.  Pronotum 
with  two  midlateral  and  usually  two  basilateral 
setae;  elytron  without  discal  setae,  with  scat- 
tered pubescence.  Elytron  without  recurrent 
groove.  Microsculpture  various.  Female  stylus 
small,  spatulate,  unisetose.  Male  parameres  as 
in  Taglianti  (1973).  Flight  wings  absent,  meta- 


thorax  markedly  reduced  in  size,  elytra  fused 
along  midline.  ABL  =  1.0-1.5  mm  in  Central 
American  species. 

NATURAL  HISTORY. — Species  of  this  genus 
live  in  deep  humus  and  decaying  leaf  litter  or 
under  deeply  embedded  stones  in  forests;  they 
can  be  found  by  the  use  of  sifting  and  Berlese 
funnel  techniques.  All  species  in  Central  Amer- 
ica are  true  alticoles;  all  are  wingless.  The  larvae 
are  unknown. 

NOTES. — The  Geocharidius  fauna  of  Middle 
America  is  very  complex  and  difficult  to  work 
with  because  of  the  lack  of  a  firm  taxonomic 
base  with  which  to  start  and  the  lack  of  repre- 
sentatives of  the  majority  of  the  fauna.  New  col- 
lecting methods  are  turning  up  these  small  hy- 
pogean  or  deep-litter  forms  in  great  numbers.  Of 
the  five  species  treated  below,  four  were  dis- 
covered as  recently  as  1974.  Use  of  these  new 
collecting  methods  and  new  tools  for  revisionary 
studies  (e.g.,  scanning  electron  micrographs) 
should  allow  detailed  studies  and  a  subsequent 
reconstruction  of  the  phylogeny  soon.  Exter- 
nally, the  beetles  differ  in  states  of  the  frontal 
furrows,  body  form,  pronotum  shape,  dorsal 
elytral  chaetotaxy  and  structure  of  the  inter- 
neurs,  and  microsculpture  even  though  the  bee- 
tles themselves  are  similar  in  habitus.  In  Central 
America,  two  habitus  types  occur:  the  de- 
pressed body  form  type  such  a  G.  integripennis 
and  the  globose  type  (Figs.  54,  55). 

Adults  can  easily  be  sexed  in  two  ways.  Males 
have  three  basal  tarsomeres  of  the  anterior  leg 
dilated  and  with  modified  setae  beneath,  forming 
a  loosely  biserrate  row,  and  two  setae  along  the 
caudal  edge  of  sternum  VI;  females  have  no  spe- 
cially modified  setae  or  dilated  tarsomeres  and 
have  four  setae  on  sternum  VI. 

ARRANGEMENT  OF  TAXA:  GEOCHARIDIUS. — 
The  Anillina  are  in  need  of  a  worldwide  review 
and  revision.  No  system  heretofore  proposed  is 
adequate  to  reflect  true  relationships.  Therefore, 
the  arrangement  of  taxa  below  is  strictly  arbi- 
trary. 

CHECKLIST  OF  GEOCHARIDIUS  SPECIES  OF 
CENTRAL  AMERICA. 

1.  G.  integripennis  (Bates  1882:145) 

2.  G.  romeoi,  new  species 

3.  G.  gimlii,  new  species 

4.  G.  phineus,  new  species 

5.  G.  tagliantii,  new  species 


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FIGURE  54.     Habitus  of  Geocharidius  phineus  n.sp.,  dorsal  aspect,  male  from  1.6  km  south  of  Pantin,  Baja  Verapaz,  Gua- 
temala. 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


487 


Key  to  Species  of  Geocharidius  of 
Central  America 

1.       Pronotum  with  lateral  edge  in  front  of 

hind  angle  multidentate  

5.  G.  tagliantii,  new  species 

-  Pronotum  with  lateral  edge  unidentate 
or  smooth  2 

2(1).  Pronotum  and  elytra  highly  convex  (lat- 
eral aspect),  especially  elytra  which  ap- 
pear to  be  'inflated'  to  a  higher  plane 
than  pronotum  3 

-  Pronotum  and  elytra  much  less  convex, 
dorsum  of  elytra  on  same  plane  with 
pronotum  (lateral  aspect)  4 

3(2).  Pronotum  with  markedly  obtuse  hind 
angle,  side  margin  in  front  of  them  not 
sinuate;  small  beetles,  SBL  =  1.3  mm 
2.  G.  romeoi,  new  species 

-  Pronotum  with  slightly  obtuse  hind  an- 
gle, side  margin  in  front  of  them  sinuate; 
larger  beetles,  SBL  =  1.5  mm 

4.  G.  phineus,  new  species 

4(2).  Pronotum  with  small  denticle  on  side 
margin  in  front  of  hind  angle;  body  long, 

narrow,  and  depressed 

1.  G.  integripennis  Bates 

-  Pronotum  without  denticle,  side  margin 
entire;  body  broader  and  less  depressed 

3.  G.  gimlii,  new  species 

1 .  Geocharidius  integripennis  (Bates) 

(Figure  73) 

Anillus  integripennis  BATES,  1882:145.  [Lectotype  d ,  GUA- 
TEMALA, Totonicapan,  Totonicapan,  14°54'N,  09P22'W 
(Champion)  (BMNH),  here  designated.] 

DIAGNOSTIC  COMBINATION. — Pronotum  with 
lateral  edge  unidentate;  pronotum  and  elytra 
slightly  convex,  dorsum  of  elytra  on  same  plane 
with  pronotum  in  lateral  aspect;  body  long,  nar- 
row, and  depressed.  Color  and  luster:  body  dark 
testaceous;  appendages  pale  testaceous.  Form: 
medium-sized  beetles  for  this  genus  with  narrow 
and  depressed  forebody  and  elytra;  pronotum 
wider  than  head  and  elytron;  eyes  absent.  Struc- 
ture: frontal  furrows  moderately  well  impressed 
anteriorly,  smooth.  Hind  angle  of  pronotum 
sharp,  with  secondary  denticle  anterior  to  it; 
side  margin  sinuate.  Elytron  with  side  margin 
narrowly  flanged  in  basal  third,  flange  microser- 


rulate,  humerus  evenly  rounded;  surface  with 
scattered  setae,  striae  absent.  Dorsal  micro- 
sculpture  of  head  and  scutellum  isodiametric, 
sculpticells  large  and  well  impressed;  effaced 
elsewhere.  Male  lectotype  not  dissected.  Size: 
see  Tables  2,  3,  and  4. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  73; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — The  type-local- 
ity is  located  in  Lower  Montane  Wet  Forest  in 
the  Guatemalan  highlands  at  about  2800  m. 

MATERIAL  EXAMINED. — Lectotype  (see  above)  and  3  para- 
lectotypes  (BMNH). 

TAXONOMIC  NOTES. — Bates  (1882)  described 
this  species  from  specimens  collected  by  Cham- 
pion. Four  of  these  were  deposited  at  BMNH 
and  labelled  with  "Biologia  Centrali-Ameri- 
cana"  labels;  another  specimen  with  the  printed 
Champion  collector  label  but  without  a  "BC-A" 
label  was  found  in  MNHP  by  Taglianti  (1973). 
Taglianti  also  found  a  specimen  with  a  hand- 
written label  indicating  it  was  from  Totonicapan, 
Guatemala.  Both  specimens  found  in  MNHP 
had  handwritten  labels  (believed  by  Taglianti  to 
be  in  Jeannel's  handwriting)  determining  the 
specimens  as  "integripennis  Bates."  Bates 
(1882: 145)  did  not  indicate  how  many  specimens 
he  studied,  but  he  provided  a  description  and  a 
handpainted  illustration  of  a  specimen  of  his  new 
species. 

Bates' s  description  and  illustration  agree  with 
the  specimens  labelled  "BC-A"  in  BMNH: 
"...  flavo-testaceus,  subnitidus;  frontalibus 
elongatis,  sat  profundis,  interspatio  subangusto 
convex."  Bates's  illustration  indicates  that  the 
specimen(s)  he  described  was  uniformly  flavo- 
testaceous  as  well. 

Unfortunately,  Taglianti  obtained  two  speci- 
mens from  MNHP  that  were  probably  studied 
by  Jeannel  and  selected  them  as  lectotype  and 
paralectotype,  and  he  used  a  page  of  text  de- 
scribing how  the  specimens  did  not  match  Jean- 
nel's 1963  redescription  of  the  species.  Taglianti 
apparently  did  not  study  the  "BC-A"  material 
at  BMNH. 

The  MNHP  specimens  are  not  Bates's  inte- 
gripennis (and  probably  are  not  what  Jeannel 
studied  earlier).  Since  the  specimens  do  not 
match  Bates's  description  (e.g.,  Taglianti  states 
that  they  have  a  darkly  colored  head  and  shal- 


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TABLE  21.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  01.  Geocharidius  romeoi  n.sp.;  map  Figure  73. 


Lat./Long. 


Elev.  Orig.  Month          No. 

(m)  deposit.          coll.          spec. 


15°14'N 
90°12'W 

1600 

USNM 

Aug 

21 

15°10'N 
90°12'W 

1500 

USNM 

Aug 

8 

Total  specimens  examined: 

29 

lowly  impressed  frontal  furrows  with  "flattened 
forehead"),  I  consider  Taglianti's  selection  of 
lectotype  null  and  void.  The  two  MNHP  speci- 
mens are  further  considered  under  G.  tagliantii, 
below. 

It  is  likely  that  Jeannel  studied  "4  femelles" 
either  from  the  BMNH  collection  (although  he 
writes  the  type  is  in  MNHP)  or  his  material  has 
been  lost.  It  is  clear  that  from  his  description 
and  drawings  he  was  describing  the  same 
species  as  Bates  (note  frontal  furrows  in  draw- 
ings and  his  statement  that  the  pronotum  "les 
cotes  faiblement  argues,  sans  sinuosite  poster- 
ieure,  non  creneles  .  ..."  It  is  no  wonder  that 
Taglianti  could  not  match  Jeannel' s  description 
with  the  specimens  at  MNHP. 

Reconstruction  of  the  probable  chain  of 
events  is  that  part  of  Champion's  material  (rep- 
resenting two  species)  went  to  BMNH,  part  to 
MNHP.  Only  the  BMNH  material  was  used  by 
Bates  in  the  original  description.  Jeannel  prob- 
ably only  studied  MNHP  material,  but  the  ma- 
terial represented  the  same  species  Bates  stud- 
ied, i.e.,  integripennis.  Perhaps  Jeannel  over- 
looked the  fact  that  he  had  two  species  rep- 
resented. Taglianti  saw  only  the  second  spe- 
cies and  misapplied  the  name. 

2.  Geocharidius  romeoi,  new  species 

(Figure  73) 

DIAGNOSTIC  COMBINATION. — Elytra  and 
pronotum  highly  convex;  pronotum  with  obtuse 
hind  angle,  and  nonsinuate  side  margin.  Color 
and  luster:  body  and  head  bright  rufotestaceous; 
appendages  pale  testaceous.  Form:  small  bee- 
tles for  this  genus  with  broad  forebody  and  in- 
flated elytra;  pronotum  highly  convex,  narrower 
than  elytra;  eyes  absent.  Structure:  frontal  fur- 
rows obsolete,  front  with  moderate-sized  tu- 
bercule  at  middle.  Hind  angle  of  pronotum  dou- 
bled, each  denticle  sharp;  side  margin  evenly 


arcuate,  not  sinuate.  Elytron  highly  convex, 
short,  side  margin  arcuate,  narrowly  flanged  in 
basal  third,  flange  microserrulate,  humerus 
evenly  rounded.  Elytron  surface  with  numerous 
scattered  setae,  striae  absent.  Dorsal  micro- 
sculpture  of  head  and  scutellum  isodiametric, 
sculpticells  large  and  well  impressed;  effaced 
elsewhere.  Male  holotype  not  dissected.  SBL  = 
1.002-1.282  mm;  TW  =  0.403-0.590  mm. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  73; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — These  beetles 
are  found  commonly  at  lower  to  middle  altitudes 
in  leaf  litter  with  Berlese  techniques.  Altitudinal 
range  extends  from  1500  m  to  1600  m  in  Tropical 
Wet  Forests.  Specimens  were  collected  in  Au- 
gust; several  were  teneral. 

TYPE-MATERIAL. — Holotype  S ,  GUATEMALA,  Baja  Vera- 
paz,  13.0  km  s  Purulha,  15°10'N,  090°12'W,  Aug.  (Ball,  Fran- 
ia,  &  Whitehead)  (USNM),  ADP  026533.  Paratvpes:  173, 
119,  same  data  as  holotype  or  as  listed  in  Table  2 1  and  de- 
posited as  USNM- 14,  UASM-4,  CAS-4,  MCZ-2,  BMNH-2, 
MNHP-2. 

DERIVATION  OF  TAXON  NAME. — Romeoi,  af- 
ter Shakespeare's  Romeo,  who  in  Act  I  told 
Benvolio,  "He  that  is  stricken  blind  cannot  for- 
get the  precious  treasure  of  his  eyesight  lost," 
in  reference  to  the  demise  of  the  Central  Amer- 
ican forests  and  its  probable  impact  on  these 
blind  beetles,  which  are  restricted  to  deep  forest 
humus  and  litter. 

3.  Geocharidius  gimlii,  new  species 
(Figure  73) 

DIAGNOSTIC  COMBINATION. — Pronotum  and 
elytra  in  lateral  aspect  on  same  plane;  posterior 
angle  of  the  pronotum  without  denticle.  Color 
and  luster:  body  and  head  flavotestaceous;  ap- 
pendages pale  testaceous.  Form:  medium-sized 
beetles  for  this  genus  with  narrow  forebody  and 
elytra;  pronotum  wider  than  head,  narrower 
than  elytron;  eyes  absent.  Structure:  frontal  fur- 
rows shallowly  impressed  anteriorly,  smooth. 
Hind  angle  of  pronotum  slightly  obtuse,  not  den- 
ticulate; side  margins  straight  and  markedly  con- 
vergent behind  middle.  Elytron  moderately  long 
and  narrow,  subdepressed,  disc  somewhat  con- 
vex; side  margin  moderately  flanged  in  basal 
third,  flange  strongly  microserrulate,  humerus 
evenly  rounded.  Elytral  surface  with  scattered 
setae,  striae  absent.  Dorsal  microsculpture  of 
large  isodiametric  meshes  on  head,  base  of 


ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


489 


FIGURE  55.     Habitus  of  Geocharidius  phineus  n.sp.,  dorsal  aspect,  male  from  1.6  km  south  of  Pantin,  Baja  Verapaz,  Gua- 
temala, SEM  composite  ca.  120x. 


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ERWIN:  GROUND-BEETLES  OF  CENTRAL  AMERICA 


491 


pronotum,  and  elytra.  Male  holotype  not  dis- 
sected. SBL  =  1.319  mm;  TW  =  0.590  mm. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  73; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — The  unique  type 
was  found  in  leaf  litter  with  Berlese  techniques 
at  an  altitude  of  2780  m  in  Lower  Montane  Wet 
Forest.  It  was  collected  in  August  and  was  not 
teneral. 

TYPE-MATERIAL. — Holotype  8,  GUATEMALA,  Huehuete- 
nango,  7.7  km  s  San  Juan  Ixcoy,  15°35'N,  091°27'W,  Aug. 
(Ball,  Frania,  &  Whitehead)  (USNM),  ADP  026556. 

DERIVATION  OF  TAXON  NAME. — Gimlii,  after 
the  dwarf  Gimli,  son  of  Gloin  (one  of  the  12 
companions  of  Thorin  Oakenshield),  who  ac- 
companied the  Hobbit,  Frodo,  on  his  trip  south, 
in  Book  I  of  The  Lord  of  the  Rings  by  Tolkien, 
in  reference  to  the  small  size  of  these  beetles. 

4.  Geocharidius  phineus,  new  species 

(Figures  54-63,  74) 

DIAGNOSTIC  COMBINATION. — Pronotum  and 
elytra  highly  convex,  elytra  inflated  to  higher 
plane  than  pronotum;  lateral  margin  of  prono- 
tum with  denticle  before  hind  angle;  front  of 
head  with  tubercule.  Color  and  luster:  body  ru- 
fotestaceous;  appendages  pale  testaceous.  Form: 
large  beetles  for  this  genus  with  highly  convex 
pronotum  and  elytra;  pronotum  very  broad, 
much  wider  than  head  and  elytron;  eyes  absent. 
Structure:  frontal  furrows  well  impressed,  front 
tuberculate  at  middle.  Hind  angle  of  pronotum 
sharp,  margin  in  front  of  angle  denticulate  and 
sinuate.  Elytron  moderately  long,  narrow,  high- 


TABLE  22.     LOCALITY  DATA  DEDUCED  FROM  SPECIMEN 
LABELS.  02.  Geocharidius  phineus  n.sp.;  map  Figure  73. 


Lat./Long. 


Elev.  Orig.  Month         No. 

(m)  deposit.          coll.          spec. 


15°16'N 
90°14'W 

1680 

USNM 

Aug 

8 

15°14'N 
90°12'W 

1600 

USNM 

Aug 

2 

15°10'N 
90°12'W 

1500 

USNM 

Aug 

1 

Total  specimens  examined: 

11 

ly  convex;  side  margin  broadly  flanged  in  basal 
third,  flange  markedly  microserrulate,  humerus 
evenly  rounded.  Elytral  surface  with  scattered 
setae,  striae  absent.  Dorsal  microsculpture  of 
large  isodiametric  meshes,  restricted  to  head, 
effaced  elsewhere.  Male  holotype  not  dissected. 
SBL  =  1.194-1.517  mm;  TW  -  0.561-0.675 
mm. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  74; 
known  only  from  the  Sierra  de  Las  Minas  of 
central  Guatemala. 

NATURAL  HISTORY  NOTES. — These  beetles 
were  collected  from  leaf  litter  with  Berlese  tech- 
niques at  altitudes  between  1500  m  and  1700  m 
in  Subtropical  Wet  Forest.  They  were  collected 
in  August;  one  was  teneral. 

TYPE-MATERIAL. — Holotype  6,  GUATEMALA,  Baja  Vera- 
paz,  1.6  km  s  Pantin,  15°16'N,  090°14'W,  Aug.  (Ball,  Frania, 
&  Whitehead)  (USNM),  ADP  026665.  Paratypes:  4<J,  6$, 
same  data  as  holotype  or  as  listed  in  Table  22  and  deposited 
as  USNM-6,  UASM-2,  CAS-2. 


FIGURES  56  to  63.  Geocharidius  phineus  n.sp.,  male  from  1.6  km  south  of  Pantin,  Baja  Verapaz,  Guatemala.  56.  Head, 
anterior  aspect,  240x.  57.  Mouthparts,  oblique  ventral  aspect,  240x.  58.  Head,  dorsal  aspect,  210x.  59.  Pronotum,  oblique 
dorsal  aspect,  210x.  60.  Prosternum,  oblique  anterior  aspect,  230x.  61.  Anterior  coxae,  left  lateral  aspect,  560x.  62.  Tibia, 
left,  anterior  aspect,  380x.  63.  Tarsomeres,  left  anterior,  oblique  ventral  aspect,  690x. 


FIGURES  64  to  69.  Geographical  distribution  maps.  64.  Bembidion  purulha  n.sp.,  squares;  B.  rogersi  Bates,  circles.  65.  B. 
mexicanum  Dejean.  66.  B.  vernale  Bates.  67.  B.  satellites  Bates,  squares;  B.  lavernae  n.sp.,  circles;  B.  quetzal  n.sp.,  triangles. 
68.  B.  diabola  n.sp.,  squares;  B.  vulcanium  Darlington,  circles;  B.  edwardsi  n.sp.,  triangles.  69.  B.  aeger  n.sp.,  squares;  B. 
chiriqui  n.sp.,  circles. 

FIGURES  70  to  74.  Geographical  distribution  maps.  70.  B.  nahuala  n.sp.,  squares;  B.  franiae  n.sp.,  circles.  71.  B.  aratum 
LeConte,  squares;  B.  cartes  n.sp.,  circles;  B.  sparsum  Bates,  triangles;  B.  armuelles  n.sp.,  arrow;  72.  B.  cyclodes  Bates, 
squares;  B.  championi  Bates,  circles;  B.  ixtatan  n.sp.,  triangles.  73.  Geocharidius  integripennis  Bates,  squares;  G.  romeoi 
n.sp.,  circles;  G.  gimlii  n.sp.,  triangles.  74.  G.  phineus  n.sp.,  squares;  G.  tagliantii  n.sp.,  circles;  B.  barrensis  n.sp.,  triangle. 


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65 


67 


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493 


74 


494 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  19 


DERIVATION  OF  NAME. — Phineus,  mythical 
prophet  struck  blind  by  Zeus,  in  reference  to  the 
lost  eyesight  of  members  of  this  species. 

5.  Geocharidius  tagliantii,  new  species 

(Figure  74) 

DIAGNOSTIC  COMBINATION. — Pronotum  in 
basal  third  with  multidentate  lateral  margins. 
Color  and  luster:  head  and  body  dark  rufous; 
appendages  pale  testaceous.  Form:  large  beetles 
for  this  genus  with  broad  forebody  and  elytra; 
pronotum  wider  than  head  and  elytron;  eyes  ab- 
sent. Structure:  frontal  furrows  not  well  im- 
pressed, rough  due  to  coarse  microsculpture. 
Hind  angle  of  pronotum  denticulate,  side  margin 
in  front  of  it  multidenticulate  and  sides  straight. 
Elytron  medium-sized,  narrow;  side  margin  nar- 
rowly flanged  in  basal  third,  flange  microserru- 
late,  humerus  squared,  side  margins  nearly  par- 
allel. Elytral  surface  with  scattered  setae,  striae 
absent.  Dorsal  microsculpture  restricted  to  head 
and  scutellum,  of  coarse  isodiametric  meshes. 
Male  aedeagus  figured  by  Taglianti  (1973). 
SBL  =  1.44-1. 47  mm. 

GEOGRAPHICAL  DISTRIBUTION. — Figure  74; 
known  only  from  the  type-locality. 

NATURAL  HISTORY  NOTES. — The  type-local- 
ity is  located  in  Lower  Montane  Wet  Forest  in 
the  Guatemalan  highlands  at  about  2800  m. 

MATERIAL  EXAMINED. — None;  this  description  is  based  on 
that  of  Taglianti  (1973). 

TYPE-MATERIAL. — Holotype  <J ,  GUATEMALA,  Totonicapan, 
Totonicapan,  14°54'N,  091°22'W  (Champion)  (MNHP). 

DERIVATION  OF  TAXON  NAME. — Named  in 
honor  of  Agusto  Vigna  Taglianti  of  Italy  who 
revised  the  Anillina  of  Mexico  and  Guatemala, 
and  who  mistakenly  regarded  this  new  species 
as  Bates's  G.  integripennis. 

ACKNOWLEDGMENTS 

I  thank  La  Verne  Jean  Magarian  for  her  fore- 
sight and  creativity  in  developing  a  computer- 
ized data  base  and  its  underlying  philosophy 
with  which  I  have  begun  production  of  a  Central 
American  faunal  study  on  ground  beetles,  this 
being  the  first  contribution  even  though  it  is  not 
part  of  the  series.  She  also  participated  in  col- 
lecting most  of  the  specimens  upon  which  this 
study  is  based.  In  addition,  I  warmly  thank  all 
those  curators  and  assistants  from  whose  mu- 
seums specimens  were  borrowed  and  which  are 
listed  under  methods. 


I  heartily  thank  George  Venable  and  Gloria 
Gordon  Zimmer  for  their  excellent  illustrative 
efforts  and  Linda  L.  Sims,  Gloria  N.  House, 
Jeannine  Weaver,  and  Noreen  Connell  for  their 
efforts  in  compiling  these  pages. 

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CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  20,  pp.  497-524,  20  figs.,  1  table 


May  14,  1982 


THE  BORNEAN  GASTROMYZONTINE  FISH 

GENERA  GASTROMYZON  AND  GLANIOPSIS 

(CYPRINIFORMES,  HOMALOPTERIDAE), 

WITH  DESCRIPTIONS  OF  NEW  SPECIES 

By 
Tyson  R.  Roberts 

California  Academy  of  Sciences,  Golden  Gate  Park, 
San  Francisco,  California  94118 


ABSTRACT:  The  endemic  Bornean  gastromyzontine  genera  Gastromyzon  and  Glaniopsis  are  revised.  Neogas- 
tromyzon is  shown  to  be  generically  distinct  from  Gastromyzon.  Gastromyzon  monticola,  formerly  placed  in  the 
synonymy  of  Gastromyzon  borneensis,  is  a  valid  species.  Five  new  species  of  Gastromyzon  (G.  contractus,  G. 
ctenocephalus,  G.  lepidogaster,  G.  megalepis,  and  G.  ridens)  and  three  new  species  of  Glaniopsis  (G.  denudata, 
G.  gossei,  and  G.  multiradiata)  are  described,  bringing  the  total  known  species  in  these  genera  to  nine  and  four, 
all  of  which  are  illustrated  photographically. 


INTRODUCTION 

Gastromyzontinae  are  bottom-dwelling  fishes 
of  swift-flowing  highland  and  lowland  streams  in 
China,  Vietnam,  and  Borneo.  Most  of  the  ap- 
proximately 15  genera  and  52  species  are  highly 
specialized,  with  ventrally  flattened  head  and 
body;  inferior  mouth  frequently  with  numerous 
barbels  or  papillae;  head,  body,  and  fins  heavily 
tuberculate;  and  paired  fins  modified.  Several 
genera  have  greatly  enlarged  pectoral  and  pelvic 
fins  with  more  numerous  rays  than  in  any  other 
Cypriniformes,  and  the  pelvic  fins  may  be  united 
posteriorly,  a  condition  otherwise  unknown  in 
the  Ostariophysi.  A  synopsis  of  all  Gastromy- 
zontinae is  given  by  Silas  (1953);  Chinese  forms 
are  reviewed  and  new  species  described  by 
Chen  (1980). 

I  collected  samples  of  Gastromyzontinae  dur- 
ing an  ichthyological  survey  of  the  Kapuas  River 
basin,  western  Borneo,  in  1976.  Work  on  their 


identification  led  to  the  present  revision  of  Gas- 
tromyzon and  Glaniopsis.  Borneo  is  inhabited 
by  five  or  six  gastromyzontine  genera:  Gastro- 
myzon Giinther,  1874;  Glaniopsis  Boulenger, 
1899;  Neogastromyzon  Popta,  1905;  Parhom- 
aloptera  Vaillant,  1902;  Progastromyzon  Hora 
and  Jayaram,  195 la,  and  Protomyzon  Hora, 
1932.  I  have  studied  only  the  first  three  of  these 
genera.  Neogastromyzon  was  placed  in  the  syn- 
onymy of  Gastromyzon  by  Inger  and  Chin 
(1961)  but  is  shown  here  to  be  a  valid  genus. 
Progastromyzon  was  placed  in  the  synonymy  of 
Protomyzon  by  Inger  and  Chin  (1962).  All  of 
these  genera,  with  the  possible  exception  of 
Protomyzon,  seem  to  be  endemic  to  Borneo. 
Two  new  species  of  Protomyzon  are  described 
from  China  by  Chen  (1980);  otherwise  Proto- 
myzon is  also  known  only  from  Borneo. 

Gastromyzon  was  revised  by  Inger  and  Chin 
(1962).  They  described  two  new  species  refera- 


[497] 


498 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  20 


ble  to  Gastromyzon,  but  misidentified  G.  bor- 
neensis,  the  generic  type-species,  and  omitted 
G.  monticola,  previously  placed  in  the  synony- 
my of  G.  borneensis  but  here  shown  to  be  a 
distinct  species.  Five  new  species  of  Gastro- 
myzon are  described  in  this  paper,  bringing  the 
total  number  of  described  species  to  nine,  four 
of  which  occur  in  the  Kapuas  basin.  Glaniopsis, 
hitherto  considered  monotypic  (Weber  and  de 
Beaufort  1916;  Inger  and  Chin  1962),  is  known 
only  from  northern  Borneo;  three  new  species 
of  Glaniopsis  are  described,  and  additional  ma- 
terial is  reported  but  not  named  which  may  rep- 
resent additional  species. 

Apart  from  systematics  little  has  been  pub- 
lished about  Gastromyzontinae.  Osteology  and 
classification  of  some  genera,  including  Gastro- 
myzon and  Glaniopsis,  are  treated  by  Ramas- 
wami  (1948,  1952);  classification  of  Glaniopsis  is 
discussed  also  by  Hora  and  Jayaram  (1951b). 
Wickler  (1971)  described  locomotion,  feeding, 
and  territorial  behavior  in  Gastromyzon ;  the  spe- 
cies he  studied  is  probably  G.fasciatus  (and  not 
G.  borneensis).  Reproductive  behavior  has  not 
been  studied. 

MATERIAL  AND  METHODS 

This  paper  is  based  on  material  deposited  in 
the  following  institutions:  British  Museum  (Nat- 
ural History)  (BMNH);  California  Academy  of 
Sciences  (CAS),  including  material  formerly  de- 
posited in  the  Natural  History  Museum,  Stan- 
ford University  (SU);  Field  Museum  of  Natural 
History,  Chicago  (FMNH);  Institut  Royal  des 
Sciences  Naturelles  de  Belgique  (IRSNB);  Mu- 
seum of  Comparative  Zoology,  Harvard  (MCZ); 
Museum  National  d'Histoire  Naturelle,  Paris 
(MNHN);  Museum  of  Zoology,  Bogor,  Indo- 
nesia (MZB);  Rijksmuseum  van  Natuurlijke  His- 
toric, Leiden  (RMNH);  and  Zoological  Mu- 
seum, University  of  Amsterdam  (ZMA). 

Specimen  lengths  are  standard  length  and  pro- 
portional measurements  are  expressed  as  times 
in  standard  length,  unless  indicated  otherwise. 
Eye  diameter  is  the  greatest  horizontal  or  nearly 
horizontal  measurement  of  the  exposed  portion 
of  the  eyeball.  Interorbital  width  is  the  mea- 
surement between  the  exposed  portion  of  the 
eyeballs.  Mental  barbels  are  referred  to  as  "flap- 
like"  rather  than  "barbel-like"  if  the  length  of 
their  base  is  greater  than  their  height  (in  some 
species  of  Glaniopsis).  "Lateral  scale  series" 


refers  to  the  scale  row  bearing  tubes  or  pores 
for  the  lateral  line  sensory  canal.  Observation  of 
scales  in  Glaniopsis  was  facilitated  by  staining 
some  specimens  superficially  with  alizarin. 
"Principal  caudal-fin  rays"  includes  all  branched 
principal  caudal  rays  plus  one  upper  and  one 
lower  simple  principal  ray. 

The  following  anatomical  features  or  charac- 
ters found  in  Gastromyzontinae  have  not  been 
named  previously,  are  relatively  unfamiliar,  or 
otherwise  require  comment: 

Adhesive  pads — pads  of  thickened,  depig- 
mented  skin  on  ventral  surface  of  anteriormost 
pectoral-  and  pelvic-fin  rays:  in  many  rheophilic, 
bottom-dwelling  cyprinoids,  probably  including 
all  Gastromyzontinae.  The  pads  apparently  in- 
crease the  frictional  properties  of  the  paired  fins 
in  contact  with  substrate;  in  Gastromyzon  con- 
tractus  they  are  covered  with  horny,  hook- 
shaped,  posteriorly  directed  projections  arising 
from  single  epidermal  cells  (Roberts,  in  press). 

Ctenoid  tubercles — tubercles  forming  minute, 
comblike  projections  on  dorsolateral  portions  of 
head:  first  reported  by  Inger  and  Chin  (1962)  as 
"numerous  short,  curved  rows  of  sensory  pa- 
pillae" on  male  Neogastromyzon  pauciradiatus ; 
I  also  found  them  on  male  Gastromyzon  cteno- 
cephalus.  In  both  species  I  observed  that  areas 
with  finely  ctenoid  tubercles  grade  into  areas 
with  more  and  more  coarsely  ctenoid  tubercles 
and  finally  into  quite  ordinary-looking  small  con- 
ical tubercles. 

Gill  opening  angular — gill  opening  lying  pos- 
terior to  opercle  and  extending  anteriorly  a  short 
distance  ventral  to  subopercle,  so  that  free  mar- 
gin of  gill  cover  is  angular:  in  some  Gastromy- 
zon. 

Gill  opening  vertical — gill  opening  lying  en- 
tirely posterior  to  opercle,  so  that  free  margin 
of  gill  cover  is  vertical  or  nearly  so:  in  some 
Gastromyzon. 

Horny  jaw  sheaths — more  or  less  thick 
sheaths  of  horny  skin,  sometimes  forming  a 
sharp  "cutting  edge,"  on  upper  and  lower  jaws: 
in  many  cyprinoids,  probably  including  all  Gas- 
tromyzontinae. 

Lateral  oral  fold — a  flap  of  skin  with  discrete 
margins,  extending  from  rostral  cap  to  pectoral- 
fin  origin  lateral  to  corner  of  mouth:  in  Gastro- 
myzon only  (Fig.  \b). 

Pastoral  pouch — an  anteriorly  open,  trans- 
verse pocket  or  pouch  on  ventral  surface  of  head 


ROBERTS:  GASTROMYZONTINE  FISH 


499 


immediately  posterior  to  mouth,  formed  by  a 
fleshy  transverse  fold  or  "postoral  flap":  in 
Gastromyzon  borneensis,  G.  monticola,  and 
Neogastromyzon  nieuwenhuisi  (Fig.  1).  Gastro- 
myzon fasciatus  sometimes  has  a  weakly  devel- 
oped postoral  flap  without  formation  of  a  post- 
oral  pouch. 

Rostral  cap — a  fold  or  cap  of  skin,  usually 
horny,  with  a  more  or  less  pronounced  ventral 
groove,  overlying  upper  lip:  in  nearly  all  cypri- 
noids,  probably  in  all  Gastromyzontinae  (Fig.  1). 
Sometimes  misidentified  as  upper  lip. 

Secondary  rostrum — a  discrete  projection  at 
snout  tip,  supported  internally  by  anteriorly  di- 
rected, flangelike  projections  of  the  lacrimal 
bones,  and  heavily  tuberculate  in  large  adults: 
in  Gastromyzon  borneensis  but  no  other  Gastro- 
myzontinae (Fig.  la,  b);  superficially  similar 
projections,  possibly  involving  the  lacrimal 
bones,  occur  in  some  Garrinae  and  Labeoinae. 

Serrae — posteriorly  or  dorsoposteriorly  di- 
rected bony  projections,  arising  from  individual 
lepidotrichia,  usually  strongly  developed  on  pel- 
vic fin,  sometimes  also  present  but  weakly  de- 
veloped on  pectoral  fin:  in  all  Gastromyzon  ex- 
cept G.  megalepis;  morphologically  similar  and 
perhaps  functionally  analogous  projections  oc- 
cur mainly  on  the  anal  fin  but  sometimes  also  on 
the  pelvic  and  other  fins  in  many  Neotropical 
Characidae,  but  they  have  not  been  reported 
previously  in  any  cyprinoids. 

Sublacrimal  groove — a  more  or  less  deep  fold 
or  groove  between  lacrimal  bone  and  rostral 
cap:  in  Gastromyzon  only  (Fig.  la,  b). 

Subopercular  groove — groove  extending  from 
lower  corner  of  gill  opening  to  pectoral-fin  ori- 
gin, sometimes  discontinuous  or  interrupted 
near  its  middle:  in  several  Gastromyzon,  possi- 
bly in  other  Gastromyzontinae  with  restricted 
gill  opening. 

Suprapelvic  flap — a  fleshy  flap  of  skin  imme- 
diately dorsal  to  anteriormost  portion  of  pelvic 
fin:  in  many  Gastromyzontinae;  in  Gastromyzon 
and  Neogastromyzon  the  suprapelvic  flap  is 
continuous  anteriorly  with  a  flat  lateral  exten- 
sion of  the  body  wall  which  forms  the  ventro- 
lateral  margin  of  the  abdomen. 

Tubercles — multicellular  horny  tubercles  of 
epidermal  origin  (frequently  referred  to  as  "nup- 
tial tubercles,"  "breeding  tubercles,"  or  "pearl 
organs"  in  the  literature)  occur  on  the  head, 
body,  and  fins  of  many  cyprinoids  including  all 


FIGURE  1.  (a)  Gastromyzon  borneensis,  sagittal  section  of 
head;  (b)  Gastromyzon  borneensis,  ventral  surface  of  head; 
(c)  Neogastromyzon  nieuwenhuisi,  ventral  surface  of  head, 
(ap  =  adhesive  pad;  b[,  b2,  b3  =  rostral,  maxillary,  mandib- 
ular  barbels;  hs  =  horny  sheaths  of  upper  and  lower  jaws;  Ib, 
Ig  =  lacrimal  bone,  sublacrimal  groove;  If  =  lateral  oral  fold; 
11,  ul  =  lower  and  upper  lips;  m  =  mandible;  pf,  pp  =  post- 
oral  fold,  postoral  pouch;  re,  rg  =  rostral  cap,  rostral  groove; 
sr  =  secondary  rostrum;  t  =  tubercle;  stippling  =  alimentary 
canal.) 


Gastromyzontinae.  They  vary  in  size  from  the 
relatively  large  ones  on  the  secondary  rostrum 
and  side  of  the  body  in  male  Gastromyzon  bor- 
neensis (Fig.  5)  to  minute  ones,  barely  visible 
with  a  dissecting  microscope,  such  as  those 
found  in  dense  patches  on  the  dorsal  surface  of 
the  pectoral-fin  rays  in  males  of  several  species 
of  Gastromyzon,  including  G.  contractus  (Fig. 
2),  or  the  very  fine  pricklelike  tubercles  widely 
scattered  on  the  head  and  body  of  Glaniopsis. 


500 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  20 


FIGURE  2.     Dense  patches  of  small,  conical  tubercles  on  dorsal  surface  of  anteriormost  pectoral-fin  rays,  Gastromyzon 
contractus,  32.2-mm  male  paratype  (CAS  44186).  Horizontal  field  width  =  4.2  mm  (a),  1.1  mm  (b),  and  330  /xm  (c). 


No  direct  observations  have  been  made  con- 
cerning their  function  in  Gastromyzontinae,  so 
a  term  which  does  not  imply  a  known  function 
is  preferred.  Tubercles  exhibit  striking  sexual 
dimorphism  in  Gastromyzontinae,  as  in  many 
other  groups  of  Cypriniformes,  but  they  are  also 
present  in  immature  specimens  of  both  sexes 
and  presumably  function  in  various  modes  of 
behavior. 

KEY  TO  BORNEAN  GENERA  OF 
GASTROMYZONTINAE 

la.  Pelvic  fins  united  posteriorly,  each  with 
14-24  branched  rays;  suprapelvic  flap 
present,  continuous  anteriorly  with  lateral 
extension  of  body  wall  which  forms  ven- 
trolateral  abdominal  margin  2 

Ib.  Pelvic  fins  separate,  each  with  7-11 
branched  rays;  suprapelvic  flap  present  or 
absent  3 

2a.  Sublacrimal  groove  present;  rostral  and 
maxillary  barbels  project  from  anterior 
margin  of  rostral  cap  (Fig.  1&);  head  width 
at  most  4;  adhesive  pad  of  anteriormost 
pectoral-fin  ray  with  anterior  margin  free 
from  abdomen;  dorsal  surface  of  pelvic 
fin  usually  serrate  Gastromyzon 

2b.  Sublacrimal  groove  absent;  rostral  and 
maxillary  barbels  project  from  posterior 
margin  of  rostral  cap;  head  width  at  least 
5;  adhesive  pad  of  anteriormost  pectoral- 
fin  ray  with  anterior  margin  continuous 
with  skin  of  abdomen;  dorsal  surface  of 
pelvic  fin  not  serrate  Neogastromyzon 

3a.  Mouth  subterminal;  nasal  barbels  pres- 


ent; two  mandibular  barbels  near  each 
corner  of  mouth  (Fig.   15);  suprapelvic 

flap  absent  Glaniopsis 

3b.  Mouth  inferior;  nasal  barbels  absent;  a 
single  mandibular  barbel  near  each  corner 
of  mouth;  suprapelvic  flap  absent?  or 
present  4 

4a.  Gill  opening  extends  ventrally  onto  ven- 
tral surface  of  body  anterior  to  pectoral 
fin;  branched  pectoral-fin  rays  16-18;  su- 
prapelvic flap  absent? Parhomaloptera 

4b.  Gill  opening  extends  ventrally  to  base  of 
pectoral  fin,  not  onto  ventral  surface  of 
body;  branched  pectoral-fin  rays  18-24; 
suprapelvic  flap  present  but  not  continued 
anteriorly  to  pelvic  fin  Protomyzon 


Gastromyzon  Giinther 

Gastromyzon  GUNTHER,  1874:454  (type-species  Gastromyzon 

borneensis  Gunther,  1874,  by  monotypy). 
Lepidoglanis  VAILLANT,  1889:81  (type-species  Lepidoglanis 

monticola  Vaillant,  1889,  by  monotypy). 

DIAGNOSIS. — Gastromyzon  differs  from  all 
other  Gastromyzontinae  including  Neogastro- 
myzon in  having  a  Sublacrimal  groove  between 
lacrimal  bone  and  rostral  cap;  rostral  and  max- 
illary barbels  projecting  from  anterior  (rather 
than  from  posterior)  margin  of  rostral  cap;  a  lat- 
eral oral  fold;  and  lepidotrichia  of  pelvic-fin  rays 
(and  to  a  lesser  extent,  of  pectoral-fin  rays)  bear- 
ing serrae  (present  in  all  but  one  species  of  Gas- 
tromyzon; absent  in  Neogastromyzon  and  not 
reported  or  observed  in  any  other  Gastromy- 
zontinae). 

Pectoral,  pelvic,  and  anal  fins  close-set  or 


ROBERTS:  GASTROMYZONTINE  FISH 


501 


even  overlapping.  Pectoral  fin  with  1  simple  and 
22-30  branched  rays,  its  origin  underneath  head 
on  or  near  a  level  with  anterior  margin  of  eye. 
Adhesive  pads  present  on  about  10-12  anterior- 
most  pectoral-fin  and  8-10  anteriormost  pelvic- 
fin  rays.  Adhesive  pad  of  first  (unbranched) 
pectoral-fin  ray  with  anterior  margin  free  from 
abdomen  (continuous  with  skin  of  abdomen  in 
Neogastromyzon).  Pelvic  fins  united  posterior- 
ly, each  with  1  simple  and  17-24  branched  rays. 
Suprapelvic  flap  large,  continuous  anteriorly 
with  lateral  extension  of  body  wall  forming  ven- 
trolateral  abdominal  margin  (as  in  Neogastro- 
myzon  but  no  other  Gastromyzontinae).  Dorsal 
fin  with  2  simple  and  7-9l/2  branched  rays.  Anal 
fin  with  2  simple  and  4-5l/2  branched  rays.  Prin- 
cipal caudal-fin  rays  usually  9+8. 

Head  and  abdomen  flattened  ventrally.  Mouth 
inferior,  transverse,  very  broad.  Nasal  barbels 
absent;  rostral  and  maxillary  barbels  small; 
mandibular  barbels  rudimentary  or  absent.  Up- 
per and  lower  jaws  with  broad,  straight-edged 
horny  jaw  sheaths.  Lower  lip  densely  papillose 
except  in  G.  ctenocephalus  (sparsely  papillose 
in  Neogastromyzon).  Gill  opening  restricted, 
lying  well  above  base  of  pectoral  fin  (as  in  Neo- 
gastromyzon  but  no  other  Bornean  Gastromy- 
zontinae). Postoral  pouch,  if  present,  with  post- 
oral  flap  originating  at  base  of  first  pectoral-fin 
ray  (postoral  pouch  originates  far  anterior  to 
pectoral-fin  origin  in  Neogastromyzon  nieuwen- 
huisi,  the  only  other  gastromyzontine  with  a 
postoral  pouch).  Specimens  over  30  mm  of  both 
sexes  usually  tuberculate,  males  generally  more 
so  than  females.  Tuberculation  heaviest  on 
snout,  dorsal  surface  of  pectoral  fin  (absent  or 
weakly  developed  on  dorsal,  anal,  and  caudal 
fins),  and  sometimes  side  of  body. 

GEOGRAPHICAL  DISTRIBUTION. — Geographi- 
cal distribution  of  Gastromyzon,  based  on  ma- 
terial examined  in  this  study,  is  illustrated  in 
Figure  3.  The  most  widely  distributed  species, 
G.  fasciatus  and  G.  lepidogaster,  occur  in 
northern,  western,  and  eastern  Borneo.  Gastro- 
myzon borneensis,  previously  reported  from 
eastern  and  western  Borneo  (Weber  and  de 
Beaufort  1916;  Inger  and  Chin  1961)  apparently 
is  restricted  to  northern  Borneo  (Sarawak  and 
North  Borneo).  Absence  of  Gastromyzon  in 
southern  Borneo  may  be  an  artifact  due  to  lack 
of  collecting;  much  of  this  region,  particularly 
the  headwaters,  is  ichthyologically  unexplored. 


Key  to  Gastromyzon 

la.  Secondary  rostrum  present  „  G.  borneensis 
Ib.  No  secondary  rostrum   2 

2a.  Sublacrimal  groove  extending  onto  side 
of  head  G.  ridens 

2b.  Sublacrimal  groove  confined  to  ventral 
surface  of  head  3 

3a.  Subopercular  groove  absent 4 

3b.  Subopercular  groove  present  6 

4a.  Postoral  pouch  present  G.  monticola 

4b.  Postoral  pouch  absent    5 

5a.  Scales  in  lateral  series  54-62;  scale  rows 
above  lateral  series  16-20;  specimens 
over  40  mm  with  abdomen  partly  or  en- 
tirely covered  with  fine  scales  

G.  lepidogaster 

5b.  Scales  in  lateral  series  44-48;  scale  rows 
above  lateral  series  11-12;  abdomen 
scaleless  except  some  specimens  with  a 
few  relatively  large  scales  between  pos- 
teriormost  pelvic-fin  rays  „  G.  megalepsis 

6a.  Snout  relatively  truncate  in  larger  speci- 
mens; dorsal  surface  of  head  and  body 
with  relatively  large,  round,  pale  spots 
(vermiculate  spots  on  head  in  small  spec- 
imens); pectoral  fin  with  thin  pale  band  at 
margin;  male  without  dense  patches  of 
small  tubercles  on  dorsal  surface  of  pec- 
toral fin  G.  punctulatus 

6b.  Snout  rounded;  dorsal  surface  of  head 
and  body  without  spots  or  with  very  small 
round  spots;  pectoral  fin  without  thin  pale 
band  at  margin;  male  with  dense  patches 
of  small  tubercles  on  dorsal  surface  of 
pectoral  fin  7 

7a.  Adpressed  dorsal  fin  extending  posterior- 
ly to  or  beyond  level  of  anal-fin  origin; 
pelvic  fin  reaching  to  or  beyond  anal-fin 
origin;  snout  sloping  downward  in  front 
of  eyes  relatively  strongly  .„.  G.  contractus 

7b.  Adpressed  dorsal  fin  falling  short  of  level 
of  anal-fin  origin;  pelvic  fin  falling  short 
of  or  just  reaching  anal-fin  origin;  snout 
sloping  downward  in  front  of  eyes  rela- 
tively gently  8 

8a.  Predorsal  scales  40-55;  branched  dorsal- 
fin  rays  8Vi;  branched  anal-fin  rays  5-5^; 
male  without  ctenoid  tubercles  on  head 
G.  fasciatus 


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110° 

I 


112° 

I 


114° 

I 


116° 

i 


118° 

i 


*  G.  borneensis 

*  G.  con  tract  us 

T  G.  ctenocephalus 
A  G.  fascist  us 

*  G.  lepidogaster 
O  G.  megalepis 


•ff  G.  monticola 
o  G.  punctulatus 
a  G.  ridens 


FIGURE  3.     Distribution  of  Gastromyzon  (based  on  material  examined). 


8b.  Predorsal  scales  about  32-36;  branched 
dorsal-fin  rays  ll/2\  branched  anal-fin  rays 
4l/2~,  male  with  ctenoid  tubercles  on  head 
G.  ctenocephalus 

Gastromyzon  borneensis  Giinther 

(Figures  la,  b,  4-5) 

Gastromyzon  borneensis  GUNTHER,  1874:6  (type-locality 
"sources  of  Mingalong  River"  =  Mengalong  River,  Beau- 
fort District,  North  Borneo). 

TYPE-MATERIAL. — The  type-series  of  five  syntypes  (BMNH 
1874.11.24:1-5)  comprises  two  species.  Four  specimens  have 
a  prominent  secondary  rostrum  and  the  abdomen  entirely 
scaleless;  they  are  conspecific.  The  largest  of  these  (BMNH 
1874.11.24:1,  79.1  mm)  is  hereby  designated  lectotype  of  G. 
borneensis;  the  other  three  (BMNH  1874.11.24:2^,  78.3, 
72.6,  and  65.5  mm)  are  paralectotypes.  The  fifth  syntype 
(BMNH  1874.11.24:5,  78.0  mm)  lacks  a  secondary  rostrum, 
has  the  abdomen  entirely  covered  with  fine  scales,  and  is  not 
conspecific  with  the  others;  it  is  now  the  holotype  of  G.  lep- 
idogaster new  species. 


ADDITIONAL  MATERIAL  EXAMINED. — NORTH  BORNEO: 
MCZ  34793,  34820,  34823,  27:27.9-70.4  mm,  Mount  Kinabalu; 
SU  31473,  ZMA  114.365,  6:32.1-52.9  mm,  Bongan  River, 
Mandu  Bay;  FMNH  68129,  4:16.5-38.4  mm,  Parutan  River, 
Tambunan;  FMNH  68128,  5:20.6-46.5  mm,  Sungei  Kainger- 
an,  Tambunan.  SARAWAK:  IRSNB  19723,  64:36.6-89.0  mm, 
Arur  Dalan,  a  torrential  tributary  of  Padapur  River,  near  Bar- 
io,  Baram  basin;  IRSNB  19724,  30:19.7-68.3  mm,  Sungai  Ra- 
mudu,  an  affluent  of  Sungai  Kalapang,  upstream  from  Para- 
mudu  village,  Baram  basin;  BMNH  1978.3.20:232-235, 
4:35.4-52.9  mm,  Medalam  River,  Gunong  Mulu;  BMNH 
1895.7.2:82,  1:65.1  mm,  Akar  River;  BMNH  1933.8.9:1-3, 
3:57.8-66.9  mm,  Lejok  River,  Tinjar  River;  FMNH  45852, 
1:70.4  mm,  Truson  River,  Lawas;  FMNH  45853,  2:64.9-70.5 
mm,  Pa  Brayong,  Truson  River;  FMNH  68583,  3:44.1-51.0 
mm,  Akah  River,  Meligong. 

DIAGNOSIS. — Attaining  89  mm,  and  thus  pos- 
sibly the  largest  species  in  the  genus,  Gastro- 
myzon borneensis  differs  from  all  of  its  conge- 
ners in  having  a  secondary  rostrum  and,  in 
sexually  mature  males,  developing  heavy  tuber- 


ROBERTS:  GASTROMYZONTINE  FISH 


503 


FIGURE  4.     Gastromyzon  borneensis,  88.2-mm  female  (IRSNB  19723). 


culation  on  posteroventral  portion  of  body.  The 
secondary  rostrum,  present  in  all  specimens  ex- 
amined over  30  mm  of  both  sexes,  is  tuberculate 
in  both  sexes,  but  becomes  slightly  larger  and 
more  tuberculate  in  males  than  in  females.  Post- 
oral  pouch  present  (absent  in  all  other  Gastro- 
myzon except  G.  monticola). 

Snout  terminating  in  secondary  rostrum,  oth- 
erwise gently  sloping  downward  in  front  of  eyes 
and  relatively  truncate.  Gill  opening  vertical,  its 
length  about  equal  to  eye  diameter;  no  suboper- 
cular  groove.  Scales  in  lateral  series  51-62;  pre- 
dorsal  scales  at  least  40;  scale  rows  above  lateral 
series  about  20;  circumpeduncular  scales  28-32. 
Pectoral  fin  falls  far  short  of  pelvic  fin,  pelvic  fin 


short  of  anal  fin,  and  depressed  dorsal  fin  far 
short  of  level  of  anal-fin  origin. 

Dorsal  and  lateral  surfaces  of  body  with  dull 
brownish  or  brownish-gray  background  broken 
by  pale,  interconnected  narrow  bands  forming 
irregularly  hexagonal  or  pentagonal  figures;  dor- 
sal surface  of  head  with  hexagonal  or  pentagonal 
figures  similar  to  those  on  body  but  much  small- 
er and  more  closely  spaced  together.  Most  col- 
oration on  fins  narrowly  confined  along  fin  rays; 
dorsal  fin  with  faintly  banded  pattern;  caudal  fin 
with  three  to  six  dusky  vertical  bands.  Paired 
fins  without  longitudinal  bands  or  pale  margins, 
pale  or  colorless  posteriorly. 


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FIGURE  5.     Gastromyzon  borneensis,  71.5-mm  male  (IRSNB  19723). 


Gastromyzon  contractus  new  species 

(Figures  2,  6) 

TYPE-MATERIAL. — The  types  and  only  known  specimens  of 
this  new  species  were  collected  in  the  mainstream  and  one 
small  tributary  of  the  Sungai  Pinoh,  a  large  high-gradient 
stream  in  the  southern  Kapuas  basin,  by  Soetikno  Woerjoat- 
modjo  and  me  in  July  1976. 

Holotype.— MZB  3447,  34.4-mm  gravid  9,  Sungai  Tebe- 
lian,  small  forest  stream  3—4  m  wide  and  50  cm  deep,  where 
it  flows  into  Sungai  Pinoh,  19  km  upstream  from  Nangapinoh. 

Paratypes. — CAS  49319,  1:30.2  mm,  same  data  as  holotype; 
MZB  3448,  CAS  44186,  USNM  230247,  RMNH  uncat., 
BMNH  uncat.,  and  MNHN  uncat.,  17:29.1-36.9  mm,  main- 
stream of  Sungai  Pinoh  20-60  km  upstream  from  Nangapinoh; 
MZB  3449  and  CAS  49320,  2:25.9-29.8  mm,  rocky  channel  in 
mainstream  of  Sungai  Pinoh  37  km  S  of  Nangapinoh. 

DIAGNOSIS. — Perhaps  the  smallest  species  of 
Gastromyzon  (largest  known  specimen  36.9 
mm),  G.  contractus  differs  from  all  of  its  con- 
geners in  having  depressed  dorsal  fin  as  well  as 
pelvic  fin  reaching  to  or  beyond  level  of  anal-fin 
origin  (depressed  dorsal  fin  not  reaching  level  of 
anal-fin  origin,  and  pelvic  fin  reaching  at  most 
to  anal  fin  in  all  other  Gastromyzon). 

Snout  sloping  downward  in  front  of  eyes  more 
strongly  than  in  most  other  Gastromyzon,  but 
not  so  strongly  as  in  G.  monticola  and  G.  ridens, 


and  broadly  rounded.  Postoral  pouch  absent. 
Gill  opening  slightly  angular,  extending  ante- 
riorly ventral  to  subopercle  only  a  short  dis- 
tance, its  length  about  equal  to  eye  diameter; 
subopercular  groove  present,  continuous  but 
shallow  or  interrupted  near  its  middle.  Scales  in 
lateral  series  52-63;  predorsal  scales  36-44; 
scale  rows  above  lateral  series  16-20;  circum- 
peduncular  scales  about  25-30.  Pectoral  fin 
overlaps  pelvic  fin.  Male  with  dense  patches  of 
small  conical  tubercles  on  dorsal  surface  of  an- 
teriormost  pectoral-fin  rays  (Fig.  2). 

Dorsal  and  lateral  surfaces  of  body  with  thick, 
pale,  interconnected  lines  forming  irregular  hex- 
agonal or  pentagonal  figures;  dorsal  surface  of 
head  with  numerous  small,  round,  dark  spots. 
Dorsal  fin  with  two  or  three  spotted  longitudinal 
bands;  caudal  fin  with  two  or  three  dark  vertical 
bands;  coloration  in  dorsal  and  caudal  fins  con- 
centrated along  fin  rays.  Paired  fins  with  faint 
markings,  pelvic  fins  sometimes  almost  entirely 
colorless  or  colorless  except  basal  portion  dus- 
ky. Some  specimens  with  dorsal  and  lateral  sur- 
faces of  body  and  head  blotchy  or  blanched, 
suggesting  physiological  or  behavioral  color 
change. 


ROBERTS:  GASTROMYZONTINE  FISH 


505 


FIGURE  6.    Gastromyzon  contractus,  34.4-mm  gravid  female  holotype  (MZB  3447). 


ETYMOLOGY. — From  the  Latin  contractus, 
drawn  together,  in  reference  to  the  overlapping 
or  nearness  of  the  fins. 

Gastromyzon  ctenocephalus  new  species 

(Figure  7) 

TYPE-MATERIAL. — This  species  is  known  only  from  the 
type-series,  collected  at  Senah,  Sarawak,  by  A.  Everett. 

Holotype.— BMNH  1893.3.6:269,  42.3-mm  <J,  Senah,  Sa- 
rawak. 

Paratypes.— BMNH  1893.3.6:261-268,  8:28.9-37.8  mm, 
same  data  as  holotype. 

DIAGNOSIS. — Gastromyzon  ctenocephalus 
differs  from  all  other  Gastromyzon  in  having 
only  8-12  papillae  on  lower  lip,  widely  spaced 
on  or  near  its  free  anterior  margin  (vs.  extremely 
numerous  papillae  densely  distributed  over  en- 
tire lower  lip);  gill  opening  strongly  angular, 
continued  farther  anteriorly  ventral  to  suboper- 
cle  than  in  any  other  Gastromyzon ;  subopercu- 
lar  groove  continuous  and  relatively  deep  (vs. 
subopercular  groove  continuous  but  relatively 


shallow,  interrupted  in  its  middle,  or  absent); 
and  sexually  mature  male  with  ctenoid  tubercles 
on  head  and  pectoral-fin  base  (ctenoid  tubercles 
otherwise  known  only  in  male  Neogastromyzon 
pauciradiatus). 

Snout  more  or  less  strongly  sloping  downward 
in  front  of  eyes,  broadly  rounded.  No  postoral 
pouch.  Length  of  gill  opening  about  1.5-2  times 
eye  diameter.  Scales  in  lateral  series  about 
50-60;  predorsal  scales  about  32-36;  scale  rows 
above  lateral  series  15-20;  circumpeduncular 
scales  28-30.  Pectoral  fin  reaching  or  slightly 
overlapping  pelvic  fin;  pelvic  fin  reaching  anal 
fin;  depressed  dorsal  fin  falling  short  of  level  of 
anal-fin  origin. 

The  holotype,  a  42.3-mm  male,  has  dense 
patches  of  small  conical  tubercles  on  dorsal  sur- 
face of  anteriormost  branched  pectoral-fin  rays, 
as  well  as  ctenoid  tubercles  on  dorsolateral  por- 
tion of  head  and  on  pectoral-fin  base.  A  32.6- 
mm  male?  has  dense  patches  of  small  tubercles 


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FIGURE  7.    Gastromyzon  ctenocephalus,  42.3-mm  ripe  male  holotype  (BMNH  1893.3.6.269). 


on  pectoral  fin  but  no  ctenoid  tubercles.  All  oth- 
er specimens,  the  three  largest  of  which, 
36.3-37.8  mm,  are  gravid  females,  lack  ctenoid 
tubercles  as  well  as  dense  patches  of  small  tu- 
bercles on  pectoral  fin,  but  have  numerous  small 
conical  tubercles  widely  scattered  on  dorsal  sur- 
face of  head  and  anteriormost  portion  of  body 
(one  tubercle  per  scale)  as  well  as  in  uniserial 
rows  of  small  conical  tubercles  on  dorsal  surface 
of  pectoral-fin  rays. 

Dorsal  surface  of  head  with  numerous  small, 
round,  pale  spots;  dorsal  and  lateral  surface  of 
body  uniformly  pale  brownish  (faded?)  or,  in 


one  paratype,  with  small,  round,  pale  spots  on 
dorsal  surface  of  body  and  pectoral-fin  base, 
similar  to  those  on  head  but  slightly  larger  and 
more  widely  spaced  apart.  In  holotype,  dorsal 
and  anal  fins  with  two  longitudinal  spotted 
bands,  spots  centered  on  rays;  caudal  fin  with 
two  or  three  faintly  spotted  vertical  bands,  and 
melanophores  narrowly  distributed  along  length 
of  all  rays;  paired  fins  with  faint  longitudinal 
bands.  In  all  paratypes,  dorsal-fin  rays  and  in- 
terradial  membranes  heavily  pigmented  except 
for  depigmented  areas  forming  an  alternating  se- 
ries of  transparent  round  spots;  anal  fin  with  in- 


ROBERTS:  GASTROMYZONTINE  FISH 


507 


FIGURE  8.     Gastromyzon  fasciatus,  59.0-mm  mature  male  (MZB  3450). 


terradial  membranes  each  bearing  a  single  ver- 
tical, oblong,  darkly  pigmented  spot  and  rays 
without  coloration;  caudal  fin  with  simple  prin- 
cipal rays  and  interradial  membranes  between 
principal  rays  unpigmented,  but  unbranched 
principal  rays  and  interradial  membranes  be- 
tween branches  of  individual  rays  darkly  pig- 
mented; paired  fins  longitudinally  banded  in 
some  paratypes,  without  noticeable  banding  in 
others;  one  paratype  with  pectoral  fin  spotted. 
Coloration  of  dorsal,  anal,  and  caudal  fins  of 
paratypes  very  different  from  that  in  other 
species  of  Gastromyzon  and  in  holotype.  This 
difference,  perhaps  due  to  individual  variation 
or  sexual  dichromatism,  should  be  investigated 
when  additional  material  becomes  available. 

ETYMOLOGY. — From  the  Greek  cteno-,  comb, 
and  cephalus,  head,  in  reference  to  the  ctenoid 
tubercles  on  the  head  of  mature  males. 

Gastromyzon  fasciatus  Inger  and  Chin 

(Figure  8) 

Gastromyzon  fasciatus  INGER  AND  CHIN,  1961:173  (type-lo- 
cality "Sungai  Dapu,  a  tributary  of  the  Baleh  River  near  the 
mouth  of  the  Sungai  Putai,  Third  Division,  Sarawak"). 

MATERIAL  EXAMINED. — SARAWAK:  FMNH  68115,  68120, 
and  BMNH  1960.12.30:1,  6:48.1-65.9  mm,  paratypes,  Baleh 


River,  Rajang  basin;  FMNH  45852,  1:70.4  mm,  Truson  River, 
Lawas;  FMNH  45853,  2:64.9-70.5  mm,  Pa  Brayong,  Truson 
River;  FMNH  68583,  3:44.1-51.0  mm,  Akah  River,  Meligong; 
SU  32378,  1:45.1  mm,  Sadong  River;  IRSNB  19725, 
6:38.6-69.2  mm,  Arur  Dalan,  torrential  tributary  of  Padapur 
River,  Baram  basin.  NORTH  BORNEO:  FMNH  68128, 
5:20.6-46.5  mm,  Sungai  Kaingeran,  Tambunan;  FMNH  un- 
cat.,  4:17.2-32.8  mm,  Parutan  River,  Tambunan.  KAPUAS  BA- 
SIN: MZB  3450  and  CAS  49321,  7:24.5-59.0  mm,  Sungai 
Pinoh,  RMNH  7636,  4:22.5-54. 1  mm,  Bongan.  MAHAKAM  BA- 
SIN: RMNH  7638,  2:53.2-56.8  mm,  B6;  RMNH  7794, 
2:39.2-54.3  mm,  Bluu  or  Bloeoe. 

DIAGNOSIS. — There  does  not  seem  to  be  any 
single  character  by  which  all  specimens  of  G. 
fasciatus  can  be  distinguished  from  all  other 
species  of  Gastromyzon,  but  it  is  nevertheless 
a  highly  distinctive  species.  Predorsal  scales 
40-55  (usually  less  than  40  in  all  other  Gastro- 
myzon). Pelvic-fin  and  anal-fin  rays  usually  more 
numerous  than  in  other  Gastromyzon  (Table  1): 
branched  pelvic-fin  rays  20-24,  modally  22  (vs. 
17-22,  modally  20  or  less);  branched  anal-fin 
rays  5-5Vi,  modally  5Y2  (vs.  4-5l/2,  modally  5  or 
less).  Specimens  of  G.  fasciatus  with  well-de- 
veloped coloration  differ  from  all  other  Gastro- 
myzon in  having  dorsal  surface  of  head  and  body 
very  darkly  pigmented,  black  or  bluish  black, 
with  narrow  vertical  white  bars  or  white  spots 


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FIGURE  9.     Gastromyzon  lepidogaster,  78.0-mm  holotype  (BMNH  1874.11.24:5). 


on  body  and  disconnected  white  vermiculations 
on  head.  This  coloration  is  well  developed  in 
freshly  preserved  specimens  from  Sarawak, 
North  Borneo,  and  the  Kapuas  basin  but  is  less 
well  developed  or  indistinct  in  older  material 
from  the  Mahakam  basin. 

Snout  gently  sloping  downward  in  front  of 
eyes,  broadly  rounded.  No  postoral  pouch.  Gill 
opening  angular,  extending  anteriorly  a  short 
distance  ventral  to  subopercle,  its  length  about 
1.5  times  eye  diameter.  Subopercular  groove 
always  present,  continuous  or  interrupted  in  the 
middle  of  its  length.  Scales  in  lateral  series 
55-68;  scales  above  lateral  series  20-23;  circum- 
peduncular  scales  30-43.  Pectoral  fin  fails  to 
reach  pelvic  fin;  pelvic  fin  falls  short  of  anal  fin; 


depressed  dorsal  fin  falls  far  short  of  level  of 
anal-fin  origin.  Male  with  dense  patches  of  small 
tubercles  on  dorsal  surface  of  anteriormost 
branched  pectoral-fin  rays. 

Freshly  preserved  specimens,  including  para- 
types  from  the  Rajang  basin  and  the  Kapuas 
material,  have  very  distinctive  coloration  differ- 
ent from  all  other  Gastromyzon :  dorsal  and  lat- 
eral surfaces  of  head  and  body  darkly  pigment- 
ed,  black  or  bluish-black,  with  several  pale  or 
white  vertical  bands  (sometimes  breaking  up 
into  a  few  large  spots)  on  body;  dorsal  fin  with 
three  or  four  longitudinal  spotted  bands,  caudal 
fin  with  two  or  three  almost  solid,  thick,  vertical 
or  irregularly  broken  black  bands;  paired  fins 
irregularly  spotted  or  mottled  (not  longitudinally 


ROBERTS:  GASTROMYZONTINE  FISH 


509 


FIGURE  10.    Gastromyzon  lepidogaster,  62.9-mm  ripe  female  paratype  (FMNH  681 13). 


banded).  Specimens  from  the  Mahakam  and 
some  other  collections,  however,  have  less  dis- 
tinctive coloration,  perhaps  partly  but  not  en- 
tirely due  to  fading  after  long  preservation. 

Gastromyzon  lepidogaster  new  species 

(Figures  9-10) 

Gastromyzon  borneensis  GUNTHER,  1874:6  (in  part);  POPTA 
1906;  WEBER  AND  DE  BEAUFORT  1916  (Fig.  1,  text;  in  part); 
INGER  AND  CHIN  1961;  1962  (in  part). 

TYPE-MATERIAL.— Holotype:  BMNH  1874.11.24:5,  78.0 
mm,  sources  of  Mengalong  River,  Beaufort  District,  North 
Borneo. 

Paratypes.— NORTH  BORNEO:  FMNH  68111-68114, 
7:38.0-64.4  mm,  Kinabatangan  basin;  FMNH  51683,  1:46.4 
mm,  East  Coast  Residency.  BRUNEI:  IRSNB  uncat.,  1:37.2 
mm,  upper  course  of  Sungai  Temburong.  KAJAN  BASIN: 
RMNH  7639,  3:79.0-83.3  mm,  upper  course  of  Boeloengan 
River  above  Ben  Barum.  MAHAKAM  BASIN:  RMNH  7635, 
5:39.3-70.1  mm,  upper  Mahakam;  RMNH  7637,  19:22.3-46.2 
mm,  Howong;  RMNH  7794,  7:36.1-83.4  mm,  Bluu  or  Bloeoe; 
RMNH  7638,  1:71.8  mm,  B6.  KAPUAS  BASIN:  RMNH  uncat., 
1:72.3  mm,  Bongan. 

DIAGNOSIS. — Gastromyzon  lepidogaster  dif- 
fers from  all  other  Gastromyzon  in  having  the 
abdomen  more  or  less  extensively  covered  with 
fine  scales  in  all  specimens  examined  over  40.2 
mm  and  in  many  below  this  size  (abdomen  en- 


tirely scaleless  in  all  other  Gastromyzon  except 
in  some  specimens  of  G.  megalepis  which  have 
a  few  relatively  large  scales  between  bases  of 
posteriormost  pelvic-fin  rays). 

Snout  gently  sloping  downward  in  front  of 
eyes,  gently  rounded.  No  postoral  pouch.  Gill 
opening  vertical,  its  length  about  equal  to  eye 
diameter.  No  subopercular  groove.  Scales  in  lat- 
eral series  54-62;  predorsal  scales  39-42;  scale 
rows  above  lateral  series  16-20;  circumpedun- 
cular  scales  30-32.  Pectoral  fin  reaches  or  falls 
short  of  pelvic  fin;  pelvic  fin  reaches  anal  fin; 
depressed  dorsal  fin  falls  short  of  level  of  anal- 
fin  origin. 

All  Gastromyzon  lepidogaster  from  North 
Borneo  and  Sarawak  have  the  abdomen  com- 
pletely covered  with  fine  scales  up  to  the  level 
of  the  anterior  margin  of  the  pectoral  girdle,  ex- 
cept the  smallest  specimen  (27.7  mm),  which  has 
the  abdomen  entirely  scaleless.  Specimens  of 
this  species  from  the  Mahakam,  Kajan,  and  Ka- 
puas  basins  above  40.2  mm,  and  many  below 
this  size,  have  the  abdomen  completely  covered 
with  fine  scales  between  the  bases  of  the  pelvic 
fins,  and  sometimes  have  scattered  scales  far- 
ther anteriorly,  but  never  have  the  abdomen 


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FIGURE  1 1.     Gastromyzon  megalepis,  54.6-mm  mature  male  holotype  (FMNH  68126). 


completely  covered  with  scales.  They  are  simi- 
lar in  other  respects  to  G.  lepidogaster  from 
North  Borneo  and  Sarawak. 

Dorsal  and  lateral  surfaces  of  head  and  body 
uniformly  brownish,  or  body  with  thin  pale  ver- 
tical bands  (similar  but  slightly  more  numerous 
than  those  in  G.  fasciatus);  bands  on  posterior 
portion  of  body  in  a  few  specimens  intercon- 
nected to  form  irregularly  pentagonal  or  hexag- 
onal figures.  Distal  half  of  dorsal  fin  with  two 
longitudinal  spotted  bands;  caudal  fin  with  broad 
dusky  vertical  band  basally  (on  portion  of  fin 
covered  by  scales)  and  three  or  four  lunate  spot- 
ted bands  on  rest  of  fin.  Anal  and  paired  fins 
with  faint  markings,  pectoral  with  spotted  lon- 
gitudinal bands,  pelvic  and  anal  with  dusky  col- 
oration largely  confined  along  rays. 

ETYMOLOGY. — From  the  Greek  lepido-,  scale, 
and  gaster,  stomach,  in  reference  to  the  exten- 
sive abdominal  squamation. 

Gastromyzon  megalepis  new  species 

(Figure  11) 

Gastromyzon  borneensis  INGER  AND  CHIN,  1961:171  (in  part). 

TYPE-MATERIAL.— Holotype:  FMNH  68126,  54.6-mm  3, 
tributary  of  Baleh  River  between  Sungai  Entunau  and  Sungai 


Putai,  Rajang  basin,  Third  District,  Sarawak,  R.  F.  Inger,  5 
Aug.  1956. 

Paratypes. — FMNH  uncut.,  5:45.1-60.5  mm,  same  data  as 
holotype;  FMNH  68127,  3:28.6-33.1  mm,  Baleh  River,  Rajang 
basin,  Third  District,  Sarawak. 

DIAGNOSIS. — Gastromyzon  megalepis  has  rel- 
atively larger  scales  than  any  other  Gastromy- 
zon ;  it  is  also  the  only  Gastromyzon  with  pelvic 
fin  lacking  serrae.  Scales  in  lateral  series  44-48 
(vs.  50  or  more  in  all  other  Gastromyzon);  pre- 
dorsal  scales  24-28  (vs.  30  or  more);  scale  rows 
above  lateral  series  11-12  (vs.  15  or  more);  and 
circumpeduncular  scales  20-21  (vs.  25  or  more). 
Pelvic-fin  rays  4-15  each  with  a  sharp,  dorso- 
posteriorly  projecting  continuous  lamina  or 
flange,  without  bony  projections  from  individual 
lepidotrichia. 

Snout  gently  sloping  downwards  in  front  of 
eyes,  broadly  rounded.  No  postoral  pouch.  Gill 
opening  vertical,  its  length  about  1.5  times  eye 
diameter.  Subopercular  groove  absent.  Pectoral 
fin  extends  to  pelvic-fin  origin;  pelvic  fin  extends 
almost  to  anal  fin;  depressed  dorsal  fin  falls  far 
short  of  level  of  anal-fin  origin.  Male  with  dense 
patches  of  minute  tubercles  on  dorsal  surface  of 
anteriormost  branched  pectoral-fin  rays. 


ROBERTS:  GASTROMYZONTINE  FISH 


511 


FIGURE  12.     Gastromyzon  monticola,  70.9-mm  ripe  female  (BMNH  1894.6.30:1%). 


Dorsal  and  lateral  surfaces  of  head  and  body 
uniformly  brownish,  sometimes  mottled  on  cau- 
dal portion  of  body.  Dorsal  fin  with  four  longi- 
tudinal spotted  bands  and  caudal  fin  with  about 
six  lunate  spotted  bands,  the  spots  nearly  con- 
fined to  fin  rays;  anal  fin  with  melanophores  uni- 
formly distributed  along  rays;  paired  fins  with 
faint  longitudinal  bands,  pale  margins. 

ETYMOLOGY. — From  the  Greek  mega-,  large, 
and  lepis,  scale,  in  reference  to  the  relatively 
large  scales  of  this  species. 


Gastromyzon  monticola  (Vaillant) 

(Figure  12) 

Lepidoglanis  monticola  VAILLANT,   1889:81   (type-locality 

"Kina-Balou"  =  Mt.  Kinabalu,  North  Borneo). 
Gastromyzon  monticola  VAILLANT,  1891;  1893:94,  pi.  1  (de- 
scription and  figures  of  syntypes). 

Gastromyzon  borneensis  WEBER  AND  DE  BEAUFORT,  1916  (in 
part). 

This  species  has  not  been  recognized  by  ich- 
thyologists since  it  was  placed  in  the  synonymy 
of  Gastromyzon  borneensis  by  Weber  and  de 


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FIGURE  13.     Gastromyzon  punctulatus,  53.0-mm  gravid  female  paratype  (FMNH  681 17). 


Beaufort;  it  was  entirely  overlooked  by  Inger 
and  Chin  (1961). 

MATERIAL  EXAMINED. — All  from  NORTH  BORNEO:  MNHN 
1889.83-85,  3:63.5-67.1  mm,  syntypes,  Mt.  Kinabalu  (Vaillant 
1889,  indicated  10  syntypes  but  only  these  3  could  be  located 
during  my  visit  to  the  MNHN  in  November  1979);  BMNH 
1894.6.30:196-197,  2:66.9-70.9  mm,  Mt.  Kinabalu  (these  two 
specimens  were  compared  directly  with  the  syntypes  during 
my  visit  to  the  MNHN);  FMNH  44725,  4:33.2-56.3  mm,  Tem- 
passuk  River,  Kota  Belud  District,  West  Coast  Residency. 

In  addition  to  the  specimens  just  listed,  I  tentatively  identify 
MCZ  54091,  2:43.1^*7.2  mm,  Sarawak,  as  this  species;  these 
specimens  were  seen  briefly  during  a  visit  to  the  MCZ  and 
have  not  been  considered  in  the  account  below. 

DIAGNOSIS. — Gastromyzon  monticola  differs 
from  all  other  Gastromyzon  except  G.  borneen- 
sis  in  having  a  postoral  pouch;  it  lacks  a  sec- 
ondary rostrum  and  differs  in  many  respects 
from  G.  borneensis.  It  differs  from  all  other 
Gastromyzon  except  G.  ridens  and  G.  contrac- 
tus  in  having  the  snout  very  strongly  sloping 
downward  in  front  of  the  eyes. 

Snout  broadly  rounded  or  slightly  pointed. 
Gill  opening  vertical,  its  length  about  equal  to 


eye  diameter.  Subopercular  groove  absent. 
Scales  in  lateral  series  about  55-58;  predorsal 
scales  about  40;  scale  rows  above  lateral  series 
20;  circumpeduncular  scales  30-32.  Pectoral  fin 
overlaps  pelvic  fin;  pelvic  fin  reaches  anal  fin; 
and  depressed  dorsal  fin  falls  far  short  of  level 
of  anal-fin  origin. 

Dorsal  and  lateral  surfaces  of  body  uniformly 
brownish;  dorsal  surface  of  head  with  intercon- 
nected thick  pale  lines  forming  irregularly  hex- 
agonal or  pentagonal  figures,  very  numerous  and 
close-set.  Dorsal,  anal,  and  caudal  fins  with  nu- 
merous small  round  spots  centered  on  rays, 
forming  about  four  longitudinal  bands  on  dorsal 
fin,  two  (faint)  bands  on  anal  fin,  and  up  to  eight 
lunate  spotted  bands  on  caudal  fin.  A  particu- 
larly prominent  dark  spot  at  dorsal-fin  origin  (a 
spot  at  dorsal-fin  origin  occurs  in  other  Gastro- 
myzon but  is  usually  not  so  noticeable).  Paired 
fins  longitudinally  banded,  with  a  moderately 
wide  pale  distal  marginal  band,  and  a  dark  sub- 
marginal  band  of  about  equal  width;  pectoral  fin 
spotted  internal  to  these  bands. 


ROBERTS:  GASTROMYZONTINE  FISH 


513 


FIGURE  14.     Gastromyzon  ridens,  50.0-mm  ripe  male  holotype  (MZB  3455). 


Gastromyzon  punctulatus  Inger  and  Chin 

(Figure  13) 

Gastromyzon  punctulatus  INGER  AND  CHIN,  1961:173  (type- 
locality  "Sungai  Dapu,  a  tributary  of  the  Baleh  River  near 
the  mouth  of  the  Sungai  Putai,  Third  Division,  Sarawak"). 

This  species  is  known  only  from  the  type- 
specimens  collected  in  tributaries  of  the  Baleh 
River,  Rajang  basin,  Sarawak. 

MATERIAL  EXAMINED.— FMNH  68117,  3:32.9-53.6  mm, 
paratypes,  same  data  as  holotype. 

DIAGNOSIS  (partly  after  Inger  and  Chin 
1961). — Larger  specimens  of  Gastromyzon 
punctulatus  differ  from  all  other  Gastromyzon 
except  G.  ridens  in  having  a  relatively  truncate 
snout  (vs.  relatively  more  rounded  in  all  other 
Gastromyzon  and  in  smaller  G.  punctulatus) 
and  a  transverse  row  of  papillae  posterior  to 
lower  lip  (not  observed  in  any  other  Gastro- 
myzon). 

Snout  gently  sloping  downward  in  front  of 
eyes.  No  postoral  pouch.  Gill  opening  slightly 


angular,  its  length  about  1.5  times  eye  diameter. 
Subopercular  groove  broadly  interrupted  or  ab- 
sent in  larger  specimens,  continuous  in  small 
specimens.  Scales  in  lateral  series  about  58-59; 
predorsal  scales  about  40-42;  scale  rows  above 
lateral  series  21-22;  circumpeduncular  scales 
about  33.  Pectoral  fin  reaches  pelvic  fin;  pelvic 
fin  falls  short  of  anal  fin;  depressed  dorsal  fin 
falls  short  of  level  of  anal-fin  origin.  Tubercles 
on  dorsal  surface  of  head,  body,  and  fins  minute 
or  absent  except  for  two  or  three  rows  of  widely 
spaced  small  conical  tubercles  on  anterior  rim 
of  snout.  Dorsal  surface  of  pectoral  fin  without 
dense  patches  of  small  tubercles. 

Two  largest  specimens  examined  by  me  (53.0- 
mm  gravid  9  and  53.7-mm  ripe  c?)  with  faint, 
pale  round  spots,  relatively  large,  all  about  the 
same  size,  and  lying  close  together,  over  entire 
dorsal  surface  of  head  and  body  including  caudal 
peduncle;  sides  of  body  otherwise  uniformly 
brownish;  paired  fins  longitudinally  banded, 
with  pale  (unpigmented)  distal  margins;  dorsal- 


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~->   —•  \o  "• 


"5      fc          a 

If  111- 

o  .3  -2  ««t: 

s    C  -S    o«  c   g 
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ROBERTS:  GASTROMYZONTINE  FISH 


515 


fin  and  anal-fin  rays  dusky,  interradial  mem- 
branes depigmented  except  for  a  few  oblong 
dark  spots;  caudal  fin  with  four  or  five  dark  nar- 
row vertical  bands.  A  small  specimen,  32.9  mm, 
is  similarly  colored,  but  spots  are  more  distinct, 
and  those  on  head  more  variable  in  shape,  ver- 
miculate  rather  than  round. 

Gastromyzon  ridens  new  species 

(Figure  14) 

TYPE-MATERIAL.— Holotype:  MZB  3455,  50.0-mm  ripe 
<$ ,  mainstream  of  Sungai  Pinoh  20-60  km  upstream  from  Nan- 
gapinoh,  Kapuas  basin,  T.  R.  Roberts,  S.  Woerjoatmodjo,  21 
July  1976. 

Paratypes.— CAS  49322,  FMNH  94228,  USNM  230249, 
BMNH  uncat.,  MNHN  uncat.,  RMNH  uncat.,  7:24.3-  35.7 
mm,  same  data  as  holotype;  BMNH  1893.3.6.270-275, 
6:35.4-48.6  mm,  Senah,  Sarawak;  SU  32378,  9:21.2-36.5  mm, 
Sadong  River,  Sarawak. 

DIAGNOSIS. — Gastromyzon  ridens  differs  from 
all  other  Gastromyzon  in  having  sublacrimal 
groove  very  deep  and  enlarged,  extending  pos- 
teriorly -onto  cheek  (vs.  sublacrimal  groove  re- 
stricted to  ventral  surface  of  head  in  all  other 
Gastromyzon);  length  of  gill  opening  less  than 
or  barely  equal  to  eye  diameter  (vs.  equal  to  or 
greater  than  eye  diameter);  and  snout  relatively 
truncate,  often  with  a  slight  ventral  concavity  at 
its  tip  (snout  more  rounded  in  all  other  Gastro- 
myzon except  G.  punctulatus,  which  lacks  ven- 
tral concavity  at  snout  tip).  Pectoral-fin  rays 
25-30,  modally  27  (vs.  22-28,  modally  26  or  less 
in  all  other  Gastromyzon;  Table  1). 

Snout  strongly  sloping  downwards  in  front  of 
eyes  (more  gently  sloping  in  all  other  Gastro- 
myzon except  G.  contractus  and  G.  monticola). 
No  postoral  pouch.  Gill  opening  vertical.  No 
subopercular  groove.  Scales  in  lateral  series 
56-78;  predorsal  scales  32-52;  scale  rows  above 
lateral  series  15-26;  circumpeduncular  scales 
about  30-40.  Pectoral  fin  considerably  overlaps 
pelvic  fin,  perhaps  more  so  than  in  any  other 
Gastromyzon  except  G.  contractus;  pelvic  fin 
falls  short  of  anal  fin;  depressed  dorsal  fin  falls 
far  short  of  level  of  anal-fin  origin.  Male  holo- 
type, 50.0  mm,  has  serrae  on  pelvic  fin  relatively 
larger  and  more  heavily  developed  than  in  any 
other  specimen  of  Gastromyzon  examined,  and 
tubercles  minute  or  absent  on  head,  body,  and 
fins  except  for  numerous  moderately  large  con- 
ical tubercles  in  a  narrow  band  near  snout  tip. 
Serration  and  tuberculation  of  paratypes  similar 
to  that  of  holotype  but  less  well  developed,  and 
some  (9  $  ?)  lack  tuberculation  on  snout  tip. 


Dorsal  and  lateral  surfaces  of  head  and  body 
entirely  or  almost  entirely  covered  with  small, 
pale  round  spots;  some  specimens  with  posterior 
portion  of  body,  body  posterior  of  head,  or  en- 
tire head  and  body  more  or  less  uniformly  pale 
or  blanched,  suggesting  behavioral  or  physiolog- 
ical color  change  (as  in  G.  contractus).  Fins 
dusky  or  faintly  marked,  median  fins  with  me- 
lanophores  mainly  confined  along  fin  rays. 

ETYMOLOGY. — From  the  Latin  ridens,  laugh- 
ing or  smiling,  in  reference  to  the  peculiar  coun- 
tenance of  this  species  caused  by  the  extension 
of  the  sublacrimal  groove  onto  the  side  of  its 
head. 

Glaniopsis  Boulenger 

Glaniopsis   BOULENGER,    1899:228  (type-species  Glaniopsis 
hanitschi  Boulenger,  1899,  by  monotypy). 


FIGURE  15.  Ventral  surface  of  head,  (a)  Glaniopsis  han- 
itschi; (b)  Glaniopsis  multiradiata  (b,,  b^,  bs  =  rostral,  max- 
illary, mental  barbels;  b3,  b4  =  mandibular  barbels;  hs  = 
horny  sheaths  of  upper  and  lower  jaws;  11,  ul  =  lower  lip, 
upper  lip;  mp  =  mental  pad). 


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110° 

I 


112° 

I 


114° 

I 


116" 

i 


118° 

I 


A  G.  hanitschi 

o  G.  multiradiata,    G.  gossei 

•  G.  denudata 
if  Membikit 

•  Baleh 
T  Tinjar 

•  Mulu 


115° 


FIGURE  16.     Distribution  of  Glaniopsis. 


DIAGNOSIS. — Glaniopsis  differs  from  all  other 
Gastromyzontinae  in  having  a  subterminal  (rath- 
er than  inferior)  mouth,  and  from  all  other  Bor- 
nean  Gastromyzontinae  in  having  two  (instead 
of  only  one)  mandibular  barbels  at  each  corner 
of  mouth  (Fig.  15). 

Dorsal  fin  with  two  simple  and  six  or  seven 
branched  rays;  anal  fin  with  two  simple  and  five 
branched  rays;  posteriormost  dorsal-fin  and 
anal-fin  rays  not  split  to  base.  Pectoral  and  pel- 
vic fins  widely  separated;  pectoral  fin  with  one 
simple  and  11-17  branched  rays,  with  an  outer 
"adhesive"  and  inner  "vibratory"  portions 
(Hora  and  Jayaram  1951),  its  origin  posterior  to 
head.  Pelvic  fins  separate,  each  with  1  simple 
and  7-10  branched  rays.  Suprapelvic  flap  ab- 
sent. Adhesive  pads  present  on  first  three  to  five 
pectoral-fin  rays  and  first  three  pelvic-fin  rays. 
Sexually  mature  males  with  small  or  minute  tu- 
bercles widely  distributed  on  head,  body,  and 
fins;  males  with  most  highly  developed  tuber- 
culation  have  median-fin  rays  and  dorsal  surface 
of  paired-fin  rays  with  uniserial  rows  of  conical 
tubercles;  tubercles  on  body  minute,  pricklelike, 
usually  one  per  scale  (many  scales  without  tu- 
bercles). Females  nontuberculate  except  for  a 
few  small  conical  tubercles  in  uniserial  rows  on 
dorsal  surface  of  anteriormost  four  or  five  pec- 
toral-fin rays. 

Key  to  Glaniopsis 

la.  Dorsal  fin  with  7  branched  rays 2 

Ib.   Dorsal  fin  with  6  branched  rays 3 


2a.   Pectoral  fin  with  15-17  and  pelvic  fin  with 

8-10  branched  rays    G.  multiradiata 

2b.  Pectoral  fin  with  12-14  and  pelvic  fin  with 

6-9  branched  rays 

Baleh,  Tinjar,  and  Mulu  Glaniopsis 

(see  accounts  below  for  additional 
information) 

3a.  Maximum  of  nine  lateral  scale  rows  (mid- 
lateral  scale  row  plus  three  scale  rows 
above  and  five  scale  rows  below  it);  prin- 
cipal caudal-fin  rays  usually  less  than  9+9 
(most  often  8+8)  G.  denudata 

3b.  At  least  20  lateral  scale  rows;  principal 
caudal-fin  rays  almost  always  9+9 4 

4a.  Caudal  peduncle  depth  7.6-9.4;  adpressed 
nasal  barbel  extends  posteriorly  at  least  to 
posterior  margin  of  eye;  eye  diameter  30- 
42,  3.5-4.6  in  interorbital  width;  horny 
sheath  of  upper  jaw  varying  from  gently 
convex  to  strongly  projecting  (beaklike) 

G.  hanitschi,  Membikit  Glaniopsis 

(see  accounts  below  for  additional 
information) 

4b.  Caudal  peduncle  depth  9.2-12.2;  adpressed 
nasal  barbel  extends  posteriorly  no  farther 
than  middle  of  eye;  eye  diameter  24-30, 
2.2-3.1  in  interorbital  width;  horny  sheath 
of  upper  jaw  varying  from  slightly  concave 
to  strongly  notched  medially  ....  G.  gossei 


ROBERTS:  GASTROMYZONTINE  FISH 


517 


FIGURE  17.     Glaniopsis  denudata,  34.5-mm  mature  male  holotype  (BMNH  1957.2.27:1). 


Glaniopsis  denudata  new  species 

(Figure  17) 

TYPE-MATERIAL.— Holotype:  BMNH  1957.2.27:1,  34.5  mm. 
Sungai  Kidikarok,  North  Borneo;  altitude  4500  ft  [ca  1370  m]; 
26  Aug.  1956;  Cambridge  North  Borneo  Expedition. 

Paratypes.— BMNH  1957.2.27:2-15,  14:19.4-34.9  mm. 
Same  collection  data  as  holotype;  IRSNB  618,  1:41.8  mm. 
Sungai  Silau  Silau,  a  headwater  tributary  of  Sungai  Liwagu, 
Labuk  basin,  near  Tenompak,  Mount  Kinabalu,  altitude  5000 
ft  [ca.  1500  m];  7  Oct.  1971;  Leopold  III,  J.  P.  Gosse. 

DIAGNOSIS. — Glaniopsis  denudata,  perhaps 
the  smallest  species  of  Glaniopsis,  differs  from 
all  other  Glaniopsis  in  having  extremely  reduced 
squamation  and  usually  fewer  than  9+9  princi- 
pal caudal-fin  rays.  Largest  specimen  a  gravid 
female,  41.8  mm.  Squamation  consists  of  lateral 
scale  series  plus  maximum  of  about  three  scale 
rows  above  and  five  below  it,  or  total  of  about 
nine  scale  rows  (vs.  20  or  more  in  all  other  Glan- 
iopsis). Principal  caudal-fin  rays  variable,  most 
often  8+8,  perhaps  always  less  than  9+9.  Sex- 
ually mature  males  with  minute,  pricklelike  tu- 
bercles scattered  on  abdomen  in  front  as  well  as 
behind  pelvic  fins.  Dorsal-fin  origin  distinctly 
posterior  to  a  vertical  through  base  of  last  pel- 
vic-fin ray.  Pelvic-fin  origin  much  nearer  pec- 
toral-fin origin  than  to  anal-fin  origin. 

Eye  17-25,  1.9-2.6  in  interorbital  space.  In- 
terorbital  space  9.7-12.7.  Branched  dorsal-fin 
rays  6(13).  Principal  caudal-fin  rays  variable, 
most  often  8+8:  ?9+?9(l),  9+8(1),  9+8  or 
9+9(1),  8+9(1),  8+8(6),  7+8(1).  Pectoral-fin 
rays  i!3(6),  i!4(6),  or  i!5(l).  Pelvic-fin  rays  i7(13). 


Length  of  nasal,  outer  mandibular,  and  some- 
times maxillary  barbels  about  equal  to  or  slightly 
more  than  eye  diameter,  otherwise  length  of  bar- 
bels less  than  eye  diameter.  Adpressed  nasal 
barbel  failing  to  reach  exposed  portion  of  eye  or 
at  most  reaching  only  slightly  beyond  front  mar- 
gin of  eye.  Mental  barbel  flaplike. 

Lateral  scale  series  incomplete,  with  102-111 
scales,  some  scales  absent  or  missing  posterior- 
ly, which,  if  present,  would  yield  counts  of 
about  1 15-120  (observations  on  three  specimens 
lightly  stained  with  alizarin).  Dorsum  and  ab- 
domen entirely  scaleless.  Tubed  scale  series 
complete  or  continuous  on  anterior  one-fourth 
to  one-half  of  body,  always  arrested  before  level 
of  dorsal-fin  origin  and  entirely  absent  poste- 
riorly. In  specimens  less  than  24  mm,  tubed 
scale  series  absent  or  absent  except  for  short 
anterior  segment  of  up  to  about  six  tube-bearing 
scales.  Myotomal  muscle  masses  more  sharply 
defined  externally  than  in  any  other  Glaniopsis, 
divisions  between  them  clearly  visible  for  entire 
length  of  body  (vs.  scarcely  noticeable  at  all,  or 
clearly  visible  only  on  posterior  third  of  body). 

All  specimens  in  type-series  with  nearly  iden- 
tical coloration.  Head  without  markings,  dusky 
dorsally  and  laterally  to  just  below  level  of  eyes, 
without  melanophores  ventrally.  Oral  barbels 
and  lips  without  melanophores.  Upper  fourth  to 
half  of  body  dusky,  with  fine  melanophores  uni- 
formly distributed  everywhere  except  in  vicinity 


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FIGURE  18.    Glaniopsis  gossei,  40.5-mm  mature  male  holotype  (IRSNB  621). 


of  dorsal-fin  base:  a  dark  area  due  to  densely 
clustered  fine  melanophores  circumscribing  dor- 
sal-fin base;  a  pallid  area  with  few  or  no  mela- 
nophores extending  on  dorsum  in  front  of  dorsal 
fin  for  a  distance  about  equal  to  dorsal-fin  base 
(some  juvenile  specimens  of  G.  hanitschi  dis- 
play a  tendency  toward  this  coloration,  but  with 
much  less  marked  contrast  between  dark  and 
pallid  areas;  similar  coloration  not  observed  in 
juveniles  or  adults  of  other  Glaniopsis).  Lower 
half  of  body  pale,  without  melanophores  except 
near  midline  of  body,  dusky  area  dorsal  to  pec- 
toral-fin base,  very  faint  dusky  area  lateral  to 
pelvic-fin  base,  and  dark  area  lateral  to  anal-fin 
base  due  to  concentrations  of  melanophores. 
Fins  unpigmented  or  dusky,  without  distinct 
markings  except  for  concentration  of  melano- 
phores on  caudal-fin  base  immediately  posterior 
to  hypural  fan. 

ETYMOLOGY. — The  feminine  adjective  denu- 
data,  Latin,  "nude,"  refers  to  the  very  limited 
squamation  and  sharply  defined  myotomal  mus- 
cles characteristic  of  the  species. 

Glaniopsis  gossei  new  species 
(Figure  18) 

TYPE-MATERIAL.— Holotype:  IRSNB  621,  40.5  mm.  Arur 
Dalan,  a  torrential  headwater  of  Sungai  Padapur,  Baram  ba- 
sin, near  Bario,  Sarawak;  altitude  3500  ft  [ca.  1 100  m];  1 1  Oct. 
1971;  Leopold  HI,  J.  P.  Gosse. 

Paratypes.— IRSNB  622,  19:28.6-46.5  mm.  Same  collection 
data  as  holotype. 


DIAGNOSIS. — Glaniopsis  gossei  differs  from 
all  other  Glaniopsis  in  having  a  series  of  depig- 
mented  vertical  bars  on  sides  of  body  (these  may 
be  lost  or  obscured  in  specimens  which  are  pal- 
lid, as  if  bleached  or  faded)  and  horny  sheath  of 
upper  jaw  strongly  notched  or  concave  medially, 
rather  than  strongly  convex  medially  (beaklike) 
or  evenly  curved.  It  also  differs  from  G.  hanit- 
schi in  having  larger  eyes,  narrower  interorbital 
space,  and  shallower  caudal  peduncle;  from  G. 
multiradiata  in  having  fewer  dorsal,  pectoral, 
and  pelvic-fin  rays,  larger  eyes,  and  longer  bar- 
bels; and  from  G.  denudata  in  having  more  ex- 
tensive squamation. 

Eye  diameter  24-30,  2.2-3. 1  in  interorbital 
space.  Interorbital  space  9.2-11.0.  Depth  caudal 
peduncle  9.2-12.2.  Branched  dorsal-fin  rays  in- 
variably 6  (20).  Principal  caudal-fin  rays  9+9  (19; 
one  specimen  with  broken  fin  not  counted).  Pec- 
toral-fin rays  ill  (2),  i!2  (14),  or  i!3  (3).  Pelvic- 
fin  rays  i7  (20).  Dorsal-fin  origin  on  a  vertical 
through  origin  of  last  pelvic-fin  ray  or  slightly 
posterior  to  it.  Pelvic-fin  origin  closer  to  anal-fin 
origin  than  to  pectoral-fin  origin. 

Length  of  nasal  and  outer  mandibular  barbels 
about  1.5  times  eye  diameter.  Length  of  other 
barbels  except  mental  barbel  about  equal  to  eye 
diameter.  Adpressed  nasal  barbel  extends  pos- 
teriorly at  least  to  middle  of  eye  but  not  beyond 
exposed  portion  of  eye.  Mental  barbel  barbel- 


ROBERTS:  GASTROMYZONTINE  FISH 


519 


FIGURE  19.    Glaniopsis  hanitschi.  (a)  59.4-mm  mature  male  (IRSNB  17540);  (b,  c)  59.4-mm  mature  female  (IRSNB  17541). 


like,  about  one-half  as  long  as  inner  mandibular 
barbel. 

Sides  of  body  completely  covered  with  scales 
which  tend  to  be  embedded  (without  free  pos- 
terior margins)  and  slightly  dispersed  (not  over- 
lapping), especially  on  posterior  half  of  body, 
making  them  difficult  to  count.  Over  100  scales 
in  lateral  series.  Dorsum  in  front  of  dorsal  fin 
scaleless,  behind  dorsal  fin  covered  with  scales. 
Entire  abdomen  scaleless(?)  except  for  a  few 
scales  near  vent  and  anal-fin  origin.  Tubed  scale 
series  complete  on  anterior  half  of  body,  varia- 
bly interrupted  on  posterior  half. 

ETYMOLOGY. — Named  for  Jean-Pierre  Gosse, 
who  collected  much  of  the  material  reported 
upon  in  this  paper,  including  the  type-series  of 
this  species. 


Glaniopsis  hanitschi  Boulenger 

(Figures  I5a,  19) 

Glaniopsis  hanitschi  BOULENGER,  1899:228  (type-locality  Ka- 
damaian  River,  Mount  Kinabalu,  altitude  2100  ft  [ca.  640 
m]). 

MATERIAL  EXAMINED.— IRSNB  17540,  52:31.7-70.8  mm, 
S  nngai  Silau  Silau,  a  headwater  tributary  of  Sungai  Liwagu, 
Labuk  basin,  near  Tenompak,  Mount  Kinabalu,  altitude  5000 
ft  [ca.  1500  m];  IRSNB  17541,  28:17.6-69.3  mm,  headwater 
tributary  of  Sungai  Liodan,  below  Bundu  Tahun,  Mount  Kin- 
abalu, altitude  4000  ft  [ca.  1200  m];  FMNH  68950,  1:53.7  mm, 
Sungai  Kelangaan,  near  Mesilan  base  camp,  Mount  Kinabalu, 
altitude  5000  ft  [ca.  1500  m];  FMNH  47985,  1:58.7  mm,  Jes- 
selton  District. 


DIAGNOSIS. — G.  hanitschi  is  the  largest  and 
most  robust  Glaniopsis,  attaining  97  mm,  with 
the  deepest  caudal  peduncle,  broadest  interor- 
bital  width,  smallest  eyes,  and  longest  barbels. 


520 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  20 


FIGURE  20.    Glaniopsis  multiradiata,  44.5-mm  female  holotype  (IRSNB  619). 


Caudal  peduncle  depth  7.6-8.9  (vs.  9.2-12.7  in 
all  other  species);  interorbital  width  7.9-9.6  (vs. 
9.2-12.7;  eye  diameter  30-42,  3.7^.6  in  inter- 
orbital  width  (vs.  17-37,  1.9-3.1  in  interorbital 
width);  and  adpressed  nasal  barbel  invariably 
extending  posteriorly  beyond  exposed  portion 
of  eye.  Mental  barbel  flaplike. 

Branched  rays  in  dorsal  fin  6  (30)  (7  in  two 
specimens  with  posteriormost  ray  abnormal),  in 
pectoral  fin  12  (20)  or  13  (12),  and  in  pelvic  fin 
6  (1),  7  (30),  or  8  (1).  Principal  caudal-fin  rays 
9+9  (22)  or  10+9  (1)  (8+9  in  one  specimen  with 
anomalous  vertebral  column). 

Sides  of  body  entirely  covered  with  close-set, 
regularly  overlapping  scales  with  exposed  pos- 
terior margins  except  for  a  small  scaleless  area 
immediately  above  base  of  pectoral  fin.  Nearly 
30  complete  scale  rows  near  middle  of  body  and 
20  or  more  on  caudal  peduncle.  Midlateral  scale 
row  complete,  with  about  125-146  scales  (19). 
Dorsal  surface  of  body  anterior  to  dorsal  fin 
varying  from  scaleless  to  covered  with  scales  for 
five-sixths  of  distance  to  occiput,  but  never 
completely  scaled.  Dorsal  surface  of  body  pos- 
terior to  dorsal  fin  completely  covered  with 
scales,  or  with  only  small  scaleless  areas  im- 
mediately posterior  to  dorsal  fin  and  on  keel-like 
ridge  over  upper  procurrent  caudal-fin  rays.  Ab- 


domen scaleless  or  nearly  scaleless  except  for 
a  few  dispersed,  embedded  scales  near  anal-fin 
base  and  just  anterior  to  vent.  Lateral  line  canal 
tube-bearing  scale  series  usually  complete  on 
anterior  half  of  body,  variably  interrupted  or  in- 
complete on  posterior  half  of  body. 

Head  without  markings.  Body  with  7-18  ver- 
tical or  slightly  oblique  dark  bars,  mainly  on  dor- 
sal half  of  sides,  partially  breaking  up  into  upper 
and  lower  portions  in  some  specimens,  and  reg- 
ularly or  irregularly  arranged,  sometimes  drop- 
ping out  or  "fusing";  some  bars  may  extend  be- 
low midlateral  line,  and  an  occasional  bar  may 
lie  entirely  below  it,  especially  in  the  caudal  re- 
gion, but  bars  do  not  continue  onto  or  across 
dorsum  either  in  front  or  behind  dorsal  fin.  An- 
teriormost  bar  usually  immediately  behind  head, 
posteriormost  just  in  front  of  caudal-fin  inser- 
tion; usually  a  bar  near  dorsal-fin  origin.  In  some 
specimens  two  or  more  bars  "fuse"  to  form  a 
single  broad  mark.  Bars  absent  or  faint  in  spec- 
imens less  than  32  mm.  A  few  larger  specimens 
with  a  narrow  darkly  pigmented  area  paralleling 
occiput  and  traversing  dorsum  immediately  pos- 
terior to  head.  Two  large  specimens  with  sepa- 
rate markings  which  resemble  bars  on  side  of 
body  and  traverse  dorsum  anterior  to  dorsal  fin. 
All  specimens  with  a  dense  cluster  of  melano- 


ROBERTS:  GASTROMYZONTINE  FISH 


521 


phores,  sometimes  faint,  on  lateral  surface  of 
pelvic-fin  base  and  sometimes  extending  onto 
basal  portion  of  first  1-2  pelvic-fin  rays.  Dorsum 
and  sides  of  body  with  lozenge-shaped  clusters 
of  melanophores  centered  on  each  scale.  Dorsal 
and  lateral  surfaces  of  head,  nasal  barbel,  upper 
lip,  dorsal  and  caudal  fins,  and  dorsal  surface  of 
paired  fins  dusky.  Ventral  and  ventrolateral  por- 
tions of  body,  ventral  surface  of  head  excluding 
upper  lip,  lower  lip,  all  barbels  except  nasal  bar- 
bel, and  ventral  surface  of  paired  fins  colorless 
or  depigmented  (without  melanophores). 

Glaniopsis  multiradiata  new  species 

(Figures  156,  20) 

TYPE-MATERIAL.— Holotype:  IRSNB  619,  44.5  mm.  Arur 
Dalan,  a  torrential  headwater  of  Sungai  Padapur,  Baram  ba- 
sin, near  Bario,  Sarawak;  altitude  3500  ft  [ca.  1 100  m];  1 1  Oct. 
1971;  Leopold  III,  J.  P.  Gosse. 

Paratypes.— IRSNB  620,  30:24.3-59.1  mm.  Same  collection 
data  as  holotype. 

DIAGNOSIS. — Glaniopsis  multiradiata  has  more 
rays  in  the  paired  fins  and  shorter  barbels  than 
any  other  Glaniopsis:  pectoral-fin  rays  J15-17, 
usually  i!6  (vs.  ill-14  in  other  species  of  Glan- 
iopsis); pelvic-fin  rays  usually  i9,  exceptionally 
i8  or  ilO  (vs.  usually  i7,  exceptionally  i6  or  i8); 
and  length  of  all  barbels  equal  to  or  less  than 
eye  diameter,  adpressed  nasal  barbel  usually 
failing  to  reach  exposed  portion  of  eye. 

Eye  diameter  20-37,  2.0-3.1  in  interorbital 
space.  Interorbital  space  9.3-12.6.  Depth  caudal 
peduncle  9.7-12.3.  Branched  dorsal-fin  rays  in- 
variably 7  (3 1).  Branched  anal-fin  rays  invariably 
5  (31).  Principal  caudal-fin  rays  9+9  (14),  9+8 
(1).  Pectoral-fin  rays  i!5  (4),  i!6  (8),  or  i!7  (7). 
Pelvic-fin  rays  i8  (2),  i9  (28),  or  i  10  (1).  Dorsal- 
fin  origin  on  a  vertical  through  origin  of  last  pel- 
vic-fin ray  or  slightly  posterior  to  it.  Pelvic-fin 
origin  closer  to  anal-fin  origin  than  to  pectoral- 
fin  origin. 

Length  of  all  barbels  usually  less  than  eye  di- 
ameter. Outer  mandibular  barbel  longest,  its 
length  usually  less  than,  but  sometimes  equal  to, 
eye  diameter.  Adpressed  nasal  barbel  usually 
failing  to  extend  posteriorly  to  exposed  portion 
of  eye,  at  most  extending  slightly  beyond  free 
anterior  margin  of  eye.  Mental  barbel  barbel- 
like,  half  as  long  as  inner  mandibular  barbel. 

Sides  of  body  completely  covered  with  close- 
set,  regularly  overlapping  scales  with  exposed 
posterior  margins,  except  for  a  small  naked  area 
just  above  pectoral-fin  base;  about  30  complete 


scale  rows  near  middle  of  body  and  20  on  caudal 
peduncle.  Lateral  scale  row  complete,  with 
about  106-130  scales  (17).  Dorsum  in  front  of 
dorsal  fin  usually  covered  with  scales  almost  to 
occiput,  always  scaled  for  at  least  half  its  length 
in  front  of  dorsal  fin.  Dorsum  posterior  to  dorsal 
fin  completely  scaled  except  for  a  small  scaleless 
area  immediately  posterior  to  dorsal  fin  in  some 
specimens.  Abdomen  posterior  to  pelvic  fins 
covered  with  embedded  scales.  Abdomen  ante- 
rior to  pelvic  fins  scaleless  except  for  a  few 
embedded  scales  median  to  pelvic-fin  bases. 
Ventral  surface  of  body  posterior  to  anal  fin 
completely  scaled.  Tubed  scale  series  either 
complete,  complete  except  for  a  few  scales  with- 
out tubes  near  end  of  series,  or  with  short  tube- 
less  segments  on  posterior  half  of  body  in  a  few 
specimens. 

Glaniopsis  multiradiata  has  a  variably  mottled 
coloration  distinct  from  that  of  other  Glaniopsis. 
Mottles  largest  and  best  defined  on  dorsum,  es- 
pecially anterior  to  dorsal  fin;  fainter  and  smaller 
but  similar  mottles  extend  onto  the  head,  in- 
cluding gill  covers,  cheeks,  and  snout  in  a  few 
specimens;  almost  all  specimens  have  mottles 
on  dorsum,  although  varying  in  distinctness,  ex- 
cept for  two  smallest  specimens,  24.3  and  29.2 
mm,  which  are  pallid,  as  if  bleached.  Majority 
of  specimens  partially  decolored,  either  on  one 
or  both  sides,  as  if  bleached.  Path  of  lateral  line 
marked  by  a  thin  dark  longitudinal  band  which 
is  absent  or  much  less  distinct  in  other  Glaniop- 
sis and  which  persists  in  specimens  otherwise 
largely  decolored.  In  specimens  with  mottlings 
on  sides  of  body,  these  tend  to  be  less  prominent 
than  those  on  dorsum.  Abdomen  colorless.  Fins 
darkly  pigmented  compared  to  other  Glaniopsis. 
Caudal  fin  in  many  specimens  with  three  wavy 
vertical  bands.  Dorsal  and  anal  fins  with  slight 
concentrations  of  melanophores  suggesting  in- 
cipient barred  pattern.  Dorsal,  anal,  and  dorsal 
surfaces  of  paired  fins  often  with  fine  melano- 
phores concentrated  along  fin  rays,  giving  them 
a  dusky  appearance.  Dorsal  surface  of  pectoral- 
fin  base  dusky  or  darkly  pigmented,  more  so 
than  in  other  Glaniopsis.  A  cluster  of  melano- 
phores on  lateral  surface  of  pelvic-fin  base. 

Compared  to  Glaniopsis  hanitschi,  G.  multi- 
radiata has  body  slightly  more  elongate  or  slen- 
der, especially  posteriorly;  snout  more  de- 
pressed; ventral  surface  of  head  flatter;  lips 
(especially  lower)  slightly  less  arcuate;  lower  lip 


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interrupted  medially  by  feebly  developed  mental 
pad  from  posterolateral  corners  of  which  mental 
barbels  arise;  bases  of  outer  and  inner  mandib- 
ular  barbels  more  widely  separated;  dorsal  fin 
slightly  smaller;  caudal  fin  (correlated  with  shal- 
lower caudal  peduncle)  much  smaller;  caudal  fin 
slightly  forked,  its  upper  lobe  slightly  larger 
(rather  than  truncate);  and  paired  fins  larger. 
Pectoral  fin  about  10-15%  longer,  its  surface 
area  when  expanded  about  one-third  greater; 
first  pectoral-fin  ray  noticeably  curved  or  bowed 
posteriorly  instead  of  nearly  straight  or  only 
slightly  curved;  fringes  on  distal  ends  of 
branched  rays  less  well  developed;  adhesive 
pads  on  ventral  surface  of  outermost  rays  thick- 
er; and  base  of  pectoral  fin  fully  one-third  long- 
er. 

ETYMOLOGY. — The  feminine  adjective  multi- 
radiata  comes  from  multus,  Latin,  "much," 
plus  radiata,  Latin,  "rayed,"  in  reference  to  the 
more  numerous  pectoral-fin  and  pelvic-fin  rays 
characteristic  of  the  species. 

Membikit  Glaniopsis 

Glaniopsis  haniischi  (in  part)  INGER  AND  CHIN,  1962: 107-108 
(specimens  from  Sungai  Membikit). 

MATERIAL  EXAMINED.— FMNH  68135,  3:46.5-52.4  mm. 
Sungai  Membikit  [tributary  of  Sungai  Pegalan,  Padas  basin, 
near  southern  end  of  Crocker  range],  Keningau  District,  Sa- 
bah,  5°22'N,  116WE. 

These  three  specimens  resemble  G.  hanitschi 
more  closely  than  they  do  any  other  Glaniopsis 
but  differ  from  typical  G.  hanitschi  from  Mount 
Kinabalu  in  having  head  narrower  and  more  de- 
pressed, fins  smaller,  and  abdomen  more  exten- 
sively covered  with  scales  anterior  to  pelvic  fins. 
Two  of  the  three  specimens  also  have  fewer  pec- 
toral-fin rays  and  a  more  incomplete  tubed  scale 
series. 

Eye  34—43,  3.5-4.1  in  interorbital  space.  In- 
terorbital  space  10.3-10.8  (vs.  7.9-9.6  in  G.  han- 
itschi). Depth  caudal  peduncle  9.0-9.4  (7.6-8.9 
in  G.  hanitschi).  Branched  dorsal-fin  rays  6  (3). 
Branched  anal-fin  rays  5  (3).  Principal  caudal-fin 
rays  9+9  (2),  9+7  (1,  with  caudal  fin  probably 
injured  or  abnormally  developed).  Pectoral-fin 
rays  ill  (2)  or  i!2  (1)  (i!2  or  more  in  G.  hanit- 
schi). Pelvic-fin  rays  i7  (3). 

Adpressed  nasal  barbel  reaching  to  or  slightly 
beyond  free  posterior  margin  of  eye.  Upper  and 
lower  horny  jaw  sheaths  markedly  convex  near 
symphysis. 

Dorsum  and  sides  of  body  nearly  completely 


covered  with  scales,  lateral  series  complete  with 
about  130-142  scales,  about  35  complete  scale 
rows  near  middle  of  body  and  28  on  caudal  pe- 
duncle. Entire  abdomen  posterior  to  pelvic  fins 
covered  with  embedded  scales;  half  of  abdomen 
anterior  to  pelvic  fins  with  embedded  scales. 
Tubed  scale  series  complete  or  nearly  complete 
on  anterior  half  of  body,  absent  or  considerably 
interrupted  on  posterior  half. 

Coloration  darker  (duskier)  than  in  G.  hanit- 
schi, without  well-defined  vertical  bars  on  a  rel- 
atively pale  background;  a  continuous  dark  mid- 
lateral  longitudinal  band  (absent  in  G.  hanitschi). 
Fine  melanophores  extending  from  sides  of  body 
further  toward  and  onto  abdomen  than  in  G. 
hanitschi.  Nearly  all  scales  with  lozenge-shaped 
cluster  of  fine  melanophores.  Largest  specimen, 
52.4  mm  female,  uniformly  dusky,  without  ver- 
tical bars  or  marks  on  dorsum  or  sides  of  body. 
Smallest  specimen,  46.5  mm  female,  with  about 
20  vertical  marks  in  side  of  body  (less  well  de- 
fined, and  lower  lying  than  vertical  bars  in  G. 
hanitschi);  dorsum  with  10  dark  transverse 
marks  anterior  and  4  posterior  to  dorsal  fin  (ab- 
sent in  G.  hanitschi).  Specimen  intermediate  in 
size,  a  well-tuberculated  47.5  mm  male,  with 
markings  on  sides  fewer  and  less  well  defined, 
but  similar  to  those  in  smallest  specimen,  and 
marks  on  dorsum  barely  visible  or  absent. 

Baleh  and  Tinjar  Glaniopsis 

MATERIAL  EXAMINED.— FMNH  68137, 68151-68153, 69868, 
24:20.4-38.0  mm,  mainstream  and  tributaries  of  Baleh  River, 
Rajang  basin,  Sarawak;  BMNH  1933.8.9.9-10,  15:19.9-39.3 
mm,  Lejok  River,  altitude  50-1500  ft  [ca.  15-460  m],  Tinjar 
River  District,  Sarawak. 

These  specimens  superficially  resemble  G. 
hanitschi  but  differ  from  it  in  their  much  smaller 
size,  shorter  barbels,  having  seven  instead  of 
only  six  branched  dorsal-fin  rays,  fewer  scales, 
and  other  minor  differences. 

Eye  24-32,  2.4-3.0  in  interorbital  width.  In- 
terorbital width  9.7-11.2.  Depth  caudal  pedun- 
cle 8.9-10.0.  Branched  dorsal-fin  rays  invariably 
7.  Principal  caudal  rays  usually  9+9.  Pectoral- 
fin  rays  i!2-i!4;  pelvic-fin  rays  i7  or  i8.  Ad- 
pressed  nasal  barbel  extends  posteriorly  to 
middle  of  eye  or  almost  to  posterior  border  of 
exposed  portion  of  eye.  Upper  and  lower  horny 
jaw  sheaths  moderately  convex  near  symphysis. 

Dorsum  and  sides  of  body  completely  covered 
with  scales,  or  scales  absent  from  dorsum  only 
a  short  distance  posterior  to  occiput;  lateral 


ROBERTS:  GASTROMYZONTINE  FISH 


523 


scale  series  with  102-115  scales,  about  18-25 
scale  rows  near  middle  of  body,  22-24  on  caudal 
peduncle.  Abdomen  between  pelvic  fins  covered 
with  scales,  otherwise  largely  scaleless. 

Dorsal  and  lateral  surfaces  of  body  mottled  or 
with  vertical  or  variably  broken  vertical  bands 
of  equal  or  variable  width.  Largest  specimen, 
39.3  mm,  a  gravid  female. 

Mulu  Glaniopsis 

MATERIAL  EXAMINED. — BMNH  uncat.,  2:18.4-33.2  mm. 
Extreme  headwater  of  Sungai  Tapin  [Baram  basin],  Gunong 
Mulu,  Sarawak. 

These  two  small  specimens  possibly  represent 
an  undescribed  species.  They  differ  from  all  oth- 
er Glaniopsis  examined  in  having  a  more  slender 
body.  Unlike  Glaniopsis  from  Membikit,  Baleh, 
and  Tinjar,  which  seem  closest  to  G.  hanitschi, 
Mulu  Glaniopsis  perhaps  is  closest  to  G.  mul- 
tiradiata.  It  agrees  with  G.  multiradiata  in  gen- 
eral appearance,  color  pattern,  and  in  having  rel- 
atively high  paired-fin  ray  counts  but  differs  in 
having  longer  barbels,  smaller  eyes,  larger 
scales,  less  extensive  squamation,  and  paired 
fins  with  slightly  fewer  rays,  as  well  as  a  more 
slender  body.  Eye  30-31  (20-25  in  5  G.  multi- 
radiata 24.3-36.9  mm),  3.4-3.7  in  interorbital 
space.  Interorbital  space  8.4-9.0.  Body  depth 
9.5-9.7  (6.4-8,1  in  G.  multiradiata).  Depth  cau- 
dal peduncle  10.2-1 1.4.  Branched  dorsal-fin  rays 
7  (2).  Branched  anal-fin  rays  5  (2).  Principal  cau- 
dal-fin rays  9+9  (2).  Pectoral-fin  rays  i!3  (1),  i!4 

(1)  (J15-17  in  G.  multiradiata).  Pelvic-fin  rays  i8 

(2)  (usually  i9  in  G.  multiradiata).  Pelvic-fin  or- 
igin closer  to  pectoral  than  to  anal-fin  origin 
(equidistant  between  pectoral  and  anal-fin 
origins  or  closer  to  anal-fin  origin  in  G.  hanitschi 
and  G.  multiradiata). 

Length  of  nasal,  rostral,  maxillary  and  outer 
mandibular  barbels  equal  to  or  greater  than  eye 
diameter.  Adpressed  nasal  barbel  reaches  at 
least  to  middle  of  eye.  Larger  specimen,  33.2 
mm,  with  mental  barbels  barbel-like.  Smaller 
specimen,  18.4  mm,  with  unusually  broad,  thin 
lower  lip  and  large  flaps  in  position  normally 
occupied  by  mental  barbels.  Lower  lip  inter- 
rupted by  a  median  mental  pad  as  in  G.  multi- 
radiata. 

Larger  specimen  with  dorsum,  abdomen,  and 
ventral  surface  of  body  posterior  to  anal  fin 
scaleless;  lateral  scale  series  nearly  complete 
with  about  100  scales,  allowing  for  4-5  scales 


missing  or  having  failed  to  develop  slightly  an- 
terior to  hypural  fan  (125-146  in  G.  hanitschi, 
106-130  in  G.  multiradiata);  at  most  6-7  scale 
rows  below  and  5  above  lateral  scale  series,  or 
a  maximum  total  of  about  13  scale  rows  on  side 
of  body  (vs.  25  or  more  in  G.  hanitschi  and  G. 
multiradiata);  tubed  scale  series  complete  on 
anterior  half  of  body,  largely  discontinuous  or 
incomplete  on  posterior  half.  Smaller  specimen 
entirely  or  almost  entirely  scaleless. 

Larger  specimen  with  dorsal  and  lateral  sur- 
faces of  head  and  body  dusky,  on  body  mainly 
due  to  large  lozenge-  or  scale-shaped  clusters  of 
melanophores  overlying  nearly  every  scale;  dor- 
sum  anterior  to  dorsal  fin  with  large  mottles  sim- 
ilar to  those  in  G.  multiradiata ;  cluster  of  me- 
lanophores lateral  to  pelvic-fin  base;  cluster  of 
melanophores  toward  base  of  each  caudal-fin 
lobe,  fins  otherwise  without  distinct  markings. 
Smaller  specimen  with  fine  melanophores  uni- 
formly distributed  over  dorsal  and  lateral  sur- 
faces of  head  and  body,  and  concentrations  of 
melanophores  along  dorsal-fin  base  and  lateral 
line  canal,  otherwise  without  distinct  markings. 

ACKNOWLEDGMENTS 

It  is  a  pleasure  to  thank  the  following  individ- 
uals who  helped  in  various  ways  during  the 
course  of  this  study:  Oliver  Crimmen,  Alwyne 
Wheeler,  Keith  Banister,  and  Gordon  Howes, 
BMNH;  Marie-Louise  Bauchot  and  Marline  De- 
soutter,  MNHN;  Marinus  Boeseman  and  Peter 
van  Helsdingen,  RMNH;  Han  Nijssen,  ZMA; 
Jean-Pierre  Gosse  and  L.  Walschaerts,  IRSNB; 
Karsten  F.  Hartel  and  William  L.  Fink,  MCZ; 
Robert  K.  Johnson,  Donald  J.  Stewart,  and 
Robert  F.  Inger,  FMNH;  and  Lillian  J.  Demp- 
ster, W.  I.  Follett,  and  Michael  E.  Hearne, 
CAS.  Photography  is  by  Alphonse  Coleman, 
MCZ;  and  Orrin  Moon,  The  Darkroom,  San  Ra- 
fael; and  scanning  electron  micrography  by 
Mary-Jacque  Mann,  SEM  lab,  National  Mu- 
seum of  Natural  History.  I  am  especially  in- 
debted to  Jean-Pierre  Gosse  for  the  opportunity 
to  report  on  the  extensive  and  beautifully  pre- 
served material  of  Gastromyzon  and  Glaniopsis 
collected  by  Leopold  III  and  himself,  and  to 
Soetikno  Woerjoatmodjo  and  Rajai  for  helping 
collect  Gastromyzon  in  the  Sungai  Pinoh. 

The  ichthyological  survey  of  the  Kapuas  basin 
was  sponsored  by  the  Museum  of  Zoology,  Bo- 
gor,  Indonesian  National  Research  Council,  and 


524 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  20 


Smithsonian  Tropical  Research  Institute.  Most 
of  the  research,  supported  by  grant  DEB77- 
24759  in  the  Systematic  Biology  Program,  Na- 
tional Science  Foundation,  was  done  at  the  Cal- 
ifornia Academy  of  Sciences  and  Tiburon  Cen- 
ter for  Environmental  Studies. 


LITERATURE  CITED 

BOULENGER,  G.  A.    1899.  Descriptions  of  two  new  homal- 

opteroid  fishes  from  Borneo.  Ann.  Mag.  Nat.  Hist. 

(7)4:228-229. 
CHEN,  Y.  1980.  Systematic  studies  on  the  fishes  of  the  family 

Homalopteridae  of  China.  II.  Classification  of  the  fishes  of 

the  subfamily  Gastromyzoninae.  [In  Chinese,  with  English 

summary.]  Acta  Hydrobiol.  Sin.  7:95-120. 
GUNTHER,  A.  1874.  Descriptions  of  new  species  of  fishes  in 

the  British  Museum.  Ann.  Mag.  Nat.  Hist.  (4)14:1-8. 
HORA,  S.  L.   1932.  Classification,  bionomics,  and  evolution 

of  homalopterid  fishes.  Mem.   Indian  Mus.  (Calcutta) 

12:263-330,  pis.  10-12. 
,  AND  K.  C.  JAYARAM.  195 la.  On  two  new  gastromy- 

zonid  fishes  from  Borneo.  Rec.  Indian  Mus.  (Calcutta) 

49:191-1%. 
,  AND .  1951b.  A  note  on  the  systematic  position 


Centra]  par  M.  le  Dr.  A.  W.  Nieuwenhuis  en  1898  et  en 
1900.  Notes  Leyden  Mus.  25:171-186. 
.  1906.  Resultats  ichthyologjques  des  voyages  scientif- 


of  the  genus  Glaniopsis  Boulenger  (Fishes:  Cyprinoidea). 

Rec.  Indian  Mus.  (Calcutta)  48:85-88. 
INGER,  R.  F.,  AND  P.  K.  CHIN.  1961.  The  Bornean  cyprinoid 

fishes  of  the  genus  Gastromyzon   Giinther.   Copeia 

1961(2):  166-176. 
,  AND .   1962.  The  fresh-water  fishes  of  North 

Borneo.  Fieldiana  Zool.  45: 1-268. 
POPTA,  C.  M.  L.   1905.  Suite  des  descriptions  preliminaires 

des  nouvelles  especes  de  poissons  recueillies  au  Borneo 


iques  de  Monsieur  le  Professeur  Dr.  A.  W.  Nieuwenhuis 
dans  le  centre  de  Borneo  (1898  et  1900).  Notes  Leyden  Mus. 
27: 1-304,  pis.  1-10. 

RAMASWAMI,  L.  S.  1948.  The  homalopterid  skull.  Proc.  Zool. 
Soc.  London  118:515-538. 

.  1952.  Skeleton  of  cyprinoid  fishes  in  relation  to  phy- 

letic  studies.  IV.  The  skull  and  other  skeletal  structures  of 
gastromyzonid  fishes.  Proc.  Natl.  Inst.  Sci.  India 
18(6):5 19-538. 

ROBERTS,  T.  R.  In  press.  Unculi  (horny  projections  arising 
from  single  cells),  an  adaptive  feature  of  the  epidermis  of 
ostariophysan  fishes.  Zool.  Scripta. 

SILAS,  E.  G.  1953.  Classification,  zoogeography  and  evolu- 
tion of  the  fishes  of  the  cyprinoid  families  Homalopteridae 
and  Gastromyzonidae.  Rec.  Indian  Mus.  (Calcutta) 
50:173-263,  pi.  5. 

VAILLANT,  L.  1889.  Sur  les  poissons  d'eaux  douces  de  Bor- 
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.  1891.  [Note  on  Lepidoglanis}.  C.  Rend.  Soc.  Philo- 
math. Paris,  1890-1891,  2:6. 

.   1893.  Contribution  a  1' etude  de  la  faune  ichthyolo- 


gique  de  Borneo.  Nouv.  Arch.  Mus.  Paris  5(3):23-l  14,  pis. 
1-2. 

-.   1902.  Resultats  zoologiques  de  1'expedition  scienti- 


fique  neerlandaise  au  Borneo  Central.  Poissons.  Notes  Ley- 
den Mus.  24:1-166,  pis.  1-2. 

WEBER,  M.,  AND  L.  F.  DE  BEAUFORT.  1916.  The  fishes  of 
the  Indo-Australian  Archipelago.  III.  Ostariophysi:  II  Cy- 
prinoidea, Apodes,  Synbranchi.  E.  J.  Brill,  Leiden, 
xv  +  455. 

WICKLER,  W.  1971.  Verhaltenstudien  am  einem  hochspezial- 
isierten  Grundfisch,  Gastromyzon  borneensis  (Cyprinoidea, 
Gastromyzonidae).  Z.  Tierpsychol.  29:467-480. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94952 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Vol.  42,  No.  21,  pp.  525-535,  33  figs.  May  14,  1982 


LATE  MIOCENE  BALANID  CIRRIPEDIA  FROM  THE  BASAL 

WILSON  RANCH  BEDS  ("MERCED"  FORMATION), 

SONOMA  COUNTY,  NORTHERN  CALIFORNIA 

By 
Victor  A.  Zullo 

Department  of  Earth  Sciences, 

University  of  North  Carolina  at  Wilmington, 

Wilmington,  North  Carolina  28403 

and 
Raj  B.  Guruswami-Naidu 

Department  of  Geology,  California  Academy  of  Sciences, 
San  Francisco,  California  94118 


ABSTRACT:  The  basal  conglomerate  of  the  Wilson  Ranch  beds  (Merced  Formation  of  authors)  contains  abun- 
dant barnacle  remains  including  three  identifiable  species  of  the  family  Balanidae.  Balanus  sp.  aff.  B.  nubilus 
Darwin,  1854,  is  similar  to  extant  B.  nubilus  and  the  California  late  Miocene-early  Pliocene  species  B.  proxi- 
nubilus  Zullo,  1979,  but  differs  in  the  internal  morphology  of  the  scutum.  Balanus  irradians  new  species  is  a 
member  of  the  Balanus  balanus  (Linnaeus,  1758)  complex  distinguished  by  its  lack  of  radii.  Notomegabalanus(?) 
insperatus  new  species  is  a  megabalanine  with  Southern  Hemisphere  cool-temperate  affinities  distinguished  by 
its  prominently  ribbed  parietes.  A  late  Miocene  age  is  assigned  to  the  basal  conglomerate  on  the  basis  of  its 
stratigraphic  relationship  to  a  radiometrically  dated  tuff  within  the  Wilson  Ranch  beds  and  on  biostratigraphic 
evaluation  of  the  molluscan  fauna.  The  fauna  of  the  basal  conglomerate  suggests  deposition  in  an  immediately 
subtidal  environment  subjected  to  wave  and  current  action,  and  a  cool  temperate  marine  hydroclimate. 

INTRODUCTION  acters  alone,  is  sufficiently  unique  to  permit  its 
A  fossiliferous  marine  conglomerate  of  late  description  as  a  new  species  with  affinities  to 
Miocene  age  in  southwestern  Sonoma  County,  the  modern  Balanus  balanus  (Linnaeus,  1758) 
northern  California  (Fig.  1)  contains  an  abun-  complex.  The  fourth  is  a  new  species  of  mega- 
dance  of  balanid  barnacle  remains.  The  speci-  balanine  that  appears  to  represent  one  of  the 
mens  consist  of  shells,  disarticulated  compart-  newlV  delimited  extant  austral  genera, 
mental  plates,  and  a  few  dissociated  scuta 

representing  four  species.  One  of  these  species  STRATIGRAPHY 
cannot  be  identified  beyond  the  generic  level.  A  Outcrops  of  marine  sandstone  in  southwest- 
second  is  similar  to,  but  not  conspecific  with,  ern  Sonoma  County  were  for  many  years  con- 
the  extant  Pacific  coast  species  Balanus  nubilus  sidered  correlatives  of  the  type  Merced  For- 
Darwin,  1854.  The  third,  based  on  shell  char-  mation  of  the  San  Francisco  peninsula  and 

[525] 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  21 


FIGURE  1.  General  locality  map  for  CASG  locality  54135, 
lower  Wilson  Ranch  beds,  southwestern  Sonoma  County,  Cal- 
ifornia. 


referred  to  that  unit  (e.g.,  Dickerson  1922; 
Weaver  1949;  Travis  1952).  Higgins  (1960)  doc- 
umented lithologic  and  faunistic  differences  be- 
tween the  type  Merced  Formation  and  the  So- 
noma deposits,  and  provided  compelling 
arguments  that  the  two  units  were  deposited  in 
separate  basins.  Higgins,  however,  did  not  pro- 
pose a  separate  formational  name  for  the  So- 
noma County  deposits. 

A  tuff  interbedded  with  the  sandstone  was 
described  by  Osmont  (1905)  as  the  Sonoma  Tuff. 
In  ensuing  years,  following  the  work  of  Dick- 
erson (1922),  the  name  Sonoma  has  been  used 
to  refer  to  the  Neogene  volcanic  assemblage  of 
eastern  Sonoma  and  Napa  counties.  The  original 
Sonoma  Tuff  can  be  regarded  as  one  of  a  series 
of  tuffaceous  intertongues  of  the  eastern  volca- 
nic assemblage  into  the  western  marine  se- 
quence. Osmont  (1905)  also  described  a  mollus- 
can  fossil  locality  in  the  marine  sandstone 
containing  the  Sonoma  Tuff,  to  which  he  applied 
the  name  Wilson  Ranch  beds.  As  this  is  the  ear- 
liest formal  stratigraphic  name  that  has  been 


used  for  this  Neogene  marine  unit,  we  will  refer 
to  these  deposits  as  the  Wilson  Ranch  beds. 

K-Ar  dates  from  the  pumiceous,  vitric  tuff  in- 
terbed  described  by  Osmont  have  given  an  age 
range  of  5.7  ±  0.6  to  6.1  ±  0.1  m.y.  (Sarna- 
Wojcicki  1976).  Bartow  and  others  (1973)  re- 
ported molluscan  assemblages  in  the  Wilson 
Ranch  beds  that  ranged  in  age  from  the  early 
Pliocene  below,  to  late  Pliocene  above  the  dated 
tuff.  The  early  and  late  Pliocene  molluscan  as- 
semblages were  correlated  with  Hemphillian 
(late  Miocene  to  early  Pliocene)  and  Blancan 
(late  Pliocene  to  early  Pleistocene)  land  mammal 
assemblages,  respectively,  from  the  Petaluma 
Formation  immediately  to  the  east.  These  rela- 
tive ages  are  in  terms  of  conventional  Pacific 
coast  usage.  If,  however,  5.0  m.y.  is  acceptable 
as  the  age  of  the  Miocene-Pliocene  boundary  as 
proposed  by  Berggren  (1972,  1978)  and  Van 
Eysinga  (1975),  then  that  part  of  the  Wilson 
Ranch  beds  below  the  dated  tuff  is  of  Miocene 
age.  This  conclusion  is  supported  by  biostrati- 
graphic  evaluation  of  the  molluscan  fauna  by 
Barry  Roth  (California  Academy  of  Sciences, 
pers.  commun.,  1981).  Roth  would  correlate  the 
lower  Wilson  Ranch  beds  with  the  Pancho  Rico 
Formation  of  Monterey  County,  California, 
which  he  and  others  now  regard  as  late  Miocene 
in  age  (Addicott  1976;  Roth  and  Gurus  wami- 
Naidu  1978). 

The  barnacles  described  herein  are  from  Cal- 
ifornia Academy  of  Sciences  Department  of  Ge- 
ology (CASG)  locality  54135.  This  locality  is  in 
a  coquina  near  the  top  of  the  basal  conglomerate 
of  the  Wilson  Ranch  beds  in  the  Two  Rock 
quadrangle,  U.S.  Geological  Survey  7.5'  series, 
1954  edition,  revised  1971,  and  is  in  that  part  of 
the  section  referred  by  Osmont  (1905)  to  the  San 
Pablo  Formation  of  the  eastern  San  Francisco 
Bay  region.  A  precise  locality  description  is  on 
file  with  the  Department  of  Geology.  At  the  ex- 
posure containing  CASG  locality  54135,  the 
Wilson  Ranch  beds  unconformably  overlie  the 
Franciscan  assemblage  (Fig.  2).  The  basal  0.5  m 
consists  of  a  fining-upwards  conglomerate  com- 
posed of  coarse  sand,  rounded  pebbles,  and  an- 
gular pebble-  to  cobble-sized  fragments  of  Fran- 
ciscan-assemblage rocks  in  a  fine-grained 
sandstone  matrix.  The  upper  part  of  the  con- 
glomerate is  the  coquina  (CASG  locality  54135) 
which  is  composed  of  predominantly  horizon- 
tally bedded  invertebrate  shells.  A  vertebrate 


ZULLO  &  GURUSWAMI-NAIDU:  NORTHERN  CALIFORNIA  MIOCENE  BALANIDS 


527 


fossil  lag,  consisting  of  scattered  intact  elements 
and  abundant,  well-rounded  bone  fragments, 
overlies  the  coquina.  The  vertebrate  lag  grades 
upward  into  3.4  m  of  massive,  moderately  in- 
durated, fine-grained  sandstone  containing  scat- 
tered invertebrates  and  rare  vertebrate  remains. 
This  sandstone  is  overlain  by  tuffaceous  sand- 
stone to  the  top  of  the  exposure.  The  basal  con- 
glomerate at  this  locality  is  estimated  to  be  50 
m  stratigraphically  below  the  dated  tuff  in  the 
Wilson  Ranch  beds  (Peter  Rodda,  California 
Academy  of  Sciences,  pers.  commun.,  1981). 

PALEOECOLOGY 

The  petrology  and  fauna  of  the  basal  conglom- 
erate suggest  that  the  barnacles  inhabited  a 
rocky  intertidal  or  immediately  subtidal  environ- 
ment exposed  to  moderate  or  heavy  surf.  The 
coquina  itself,  and  its  relationship  to  the  under- 
lying fining-upwards  conglomerate,  suggest  a 
time  of  stillstand  after  rapid  transgression  in  a 
shallow  depositional  basin  affected  by  wave 
base  and/or  current  action.  According  to  Barry 
Roth  (pers.  commun.,  1981)  the  molluscan  fauna 
suggests  a  marine  hydroclimate  similar  to  that 
of  modern  Puget  Sound  (i.e.,  cool  temperate). 
These  environmental  conditions  are  supported 
by  the  composition,  morphology,  and  preser- 
vation of  the  barnacle  fauna. 


FIGURE  2.  Stratigraphic  setting  for  CASG  locality  54135, 
lower  Wilson  Ranch  beds,  southwestern  Sonoma  County,  Cal- 
ifornia. 


SYSTEMATIC  ACCOUNT 

Subclass  CIRRIPEDIA  Burmeister,  1834 

Order  THORACICA  Darwin,  1854 
Superfamily  BALANOIDEA  (Darwin),  Newman 

and  Ross,  1976 

Family  BALANIDAE  Darwin,  1854 
Subfamily  BALANINAE  Darwin,  1854 

Balanus  sp.  aff.  B.  nubilus  Darwin,  1854 
(Figures  3-11) 

Four  complete  shells,  one  partial  shell,  sev- 
eral disarticulated  compartmental  plates,  one 
entire  scutum,  four  fragmentary  scuta,  and  a  few 
fragments  of  basis  from  CASG  locality  54135  are 
here  referred  to  a  species  that  is  similar  to,  but 
probably  not  conspecific  with,  Balanus  nubilus. 
The  shells  are  conic,  the  radii  are  moderately 
well  developed  with  oblique,  jagged  summits, 
and  the  orifice  is  toothed.  The  ribbing  of  the 
shell  interior  is  not  prominent,  and  the  basis  in 
larger  individuals  is  cup-shaped  and  profusely 
porous  near  the  margins.  The  parietal  septa  are 


crowded  and  thin,  and  the  narrow,  elongate  pa- 
rietal tubes  rarely  possess  transverse  septa. 

The  scuta  that  appear  to  be  associated  with 
these  shells  are  higher  than  wide,  slightly  bowed 
outwards  near  the  apex,  and  with  a  slightly  re- 
flexed  tergal  margin.  The  exterior  is  ornamented 
by  prominent,  closely  spaced  growth  ridges  that 
are  faintly  crenate,  and  the  basitergal  angle  is 
rounded.  The  articular  ridge  is  short,  less  than 
one-half  the  length  of  the  tergal  margin,  and 
slightly  reflexed  over  the  narrow,  shallow,  artic- 
ular furrow.  The  adductor  ridge  is  erect,  highest 
along  the  margin  of  the  large,  oval,  and  deeply 
impressed  adductor  muscle  pit,  but  extending 
both  apically  and  basally  beyond  the  boundaries 
of  the  adductor  pit.  The  depressor  muscle  pit  is 
large,  triangular,  shallow,  and  bordered  on  its 
occludent  margin  by  a  low  ridge.  The  depressor 
muscle  pit  has  one  or  more  narrow,  low  longi- 
tudinal ridges  within.  The  depressor  muscle  pit 
located  at  the  basioccludent  angle  is  triangularly 


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;^ 


FIGURES  3-11.  Balanus  sp.  aff.  5.  nubilus  Darwin,  1854.  Figs.  3-5.  Exterior,  interior,  and  basal  views  of  lateral  plate, 
hypotype  CASG  60881 ;  greatest  height  47.5  mm.  Figs.  6-7.  Basal  and  alar  sutural  edge  views  of  carinorostral  and  carinal  plates, 
hypotype  CASG  60882;  greatest  height  61  mm,  greatest  wall  thickness  17  mm  (note  transverse  septa  in  parietal  tubes).  Figs. 
8-9.  Interior  and  exterior  views  of  scutum,  hypotype  CASG  60883;  height  23  mm.  Fig.  10.  Side  view  of  shell,  hypotype  CASG 
60884;  greatest  height  49  mm.  Fig.  11.  Fragment  of  basis,  hypotype  CASG  60885;  greatest  diameter  of  fragment  35.5  mm. 


ZULLO  &  GURUSWAMI-NAIDU:  NORTHERN  CALIFORNIA  MIOCENE  BALANIDS 


529 


elongate,  deeply  impressed,  and  partially  over- 
hung by  the  occludent  margin  of  the  plate. 

The  shell  of  this  species  is  similar  to  that  of 
Balanus  nubilus  and  the  related  fossil  species  B. 
proxinubilus  Zullo,  1979,  from  the  upper  Mio- 
cene Pancho  Rico  Formation  of  central  Califor- 
nia and  the  upper  Pliocene  San  Diego  Formation 
of  southern  California.  The  only  qualitative  dif- 
ference is  that  the  radii  of  the  Wilson  Ranch 
species  appear  to  be  consistently  broader  than 
those  of  either  B.  nubilus  or  B.  proxinubilus. 
The  Wilson  Ranch  scuta  differ  in  having  the  ad- 
ductor ridge  separate  from  the  articular  ridge, 
and  in  the  much  shorter  articular  ridge.  In  this 
regard  the  internal  morphology  is  somewhat 
similar  to  that  of  the  scutum  of  B.  rostratus 
Hoek,  1883,  but  the  presence  of  vertical  ridges 
or  crests  in  the  depressor  muscle  pit  serves  to 
distinguish  the  Wilson  Ranch  scuta  and  to  sug- 
gest relationship  with  B.  nubilus. 

Balanus  irradians  new  species 
(Figures  12-15,  18-21) 

DIAGNOSIS. — Shell  of  six  compartmental 
plates  without  radii;  alae  restricted  to  sheath 
area;  sutures  between  compartmental  plates  ob- 
scured or  represented  by  linear  grooves;  pari- 
etes  with  large,  square  to  rectangular,  trans- 
versely septate  parietal  tubes;  number  of  interior 
ribs  greater  than  number  of  parietal  septa;  basis 
calcareous,  solid;  distinguished  from  other 
members  of  the  Balanus  balanus  complex  by 
the  lack  of  radii. 

DESCRIPTION. — Shell  thick,  low  to  high  conic 
or  dome-shaped,  with  narrow  carinolaterals  and 
a  small,  untoothed,  diamond-shaped  orifice;  su- 
tures between  the  six  compartmental  plates  ob- 
scured or  represented  by  narrow  grooves;  radii 
absent;  sutural  edges  between  adjacent  com- 
partmental plates  broad,  bearing  coarse,  com- 
plexly arborescent  denticulations;  alae  narrow, 
with  horizontal  summits,  and  restricted  to  region 
of  sheath;  exterior  of  parietes  usually  corroded; 
uncorroded  parietes  with  external  ornamenta- 
tion of  irregular,  transverse  growth  rugae 
crossed  by  fine  longitudinal  striae;  some  speci- 
mens bear  one  to  three  external  ribs  on  larger 
compartmental  plates;  length  of  sheath  at  least 
one-half  height  of  compartmental  plate;  lower 
edge  of  sheath  free-standing,  acute,  with  cavity 
between  it  and  interior  of  shell  wall;  interior  of 
shell  wall  below  sheath  strongly  ribbed,  with  the 


largest  internal  ribs  corresponding  to  the  parietal 
septa,  and  from  one  to  three  smaller  ribs  be- 
tween parietal  septa;  parietal  tubes  large,  nearly 
square  to  rectangular,  crossed  by  transverse 
septa  and  sometimes  secondarily  filled  in  upper 
third;  parietal  septa  denticulate  basally;  inner 
surface  of  external  lamina  bears  denticulae  be- 
tween parietal  septa;  basis  calcareous,  solid. 

MATERIAL  EXAMINED. — Twelve  whole  shells  and  26  disar- 
ticulated compartmental  plates  from  CASG  locality  54135. 

TYPE  MATERIAL.— Holotype,  CASG  no.  60891;  paratypes, 
CASG  nos.  60892  through  60909,  and  paratype  lot  CASG  no. 
60910  in  the  California  Academy  of  Sciences  paleontological 
type  collection. 

ETYMOLOGY. — The  specific  name  is  derived 
from  the  Latin  prefix  />-,  without,  and  radius. 

DISCUSSION. — Balanus  irradians  and  the  ex- 
tant boreo-arctic  species  B.  balanus  and  B.  ros- 
tratus, constitute  a  small  group  in  the  genus  Bal- 
anus that  is  characterized  by  solid,  calcareous 
bases,  large,  square,  parietal  tubes,  and  second- 
ary ribs  between  the  primary  septal  ribs  of  the 
interior  lamella  of  the  parietal  wall.  The  lack  of 
radii  and  the  overall  morphology  associated  with 
a  shell  lacking  radii  are  the  major  features  dis- 
tinguishing B.  irradians  from  B.  balanus  and  B. 
rostratus.  In  other  respects,  B.  irradians  shares 
characteristics  with  the  extant  species.  The 
prominent  external  ribs  present  on  some  speci- 
mens are  reminiscent  of  the  ribbed  wall  of  B. 
balanus,  rather  than  the  smooth  shell  of  B.  ros- 
tratus. On  the  other  hand,  both  B.  irradians  and 
B.  rostratus  have  transverse  parietal  septa  that 
are  lacking  in  B.  balanus. 

The  corroded  nature  of  the  shells  and  the  type 
of  sediment  in  which  they  occur  suggest  that  B. 
irradians  lived  in  moderate-  to  high-energy  en- 
vironments. The  thick  shell  wall,  the  suppres- 
sion of  radii,  and  the  consequent  development 
of  a  broad,  coarsely  denticulate  sutural  area  for 
rigid  articulation  of  the  compartmental  plates  are 
comparable  to  shell  development  in  species  of 
Tetraclita  Schumacher,  1817,  that  often  inhabit 
areas  of  moderate  to  heavy  wave  shock  and 
abrasion  in  the  intertidal  zone.  A  consequence 
of  the  lack  of  radii  is  that  the  orifice  cannot  be 
enlarged  by  lateral  growth  along  the  sutural 
edges  of  the  plates  as  the  shell  increases  in  basal 
diameter  and  height.  As  Ross  (1969)  showed  for 
Tetraclita,  monometric  growth  forms  enlarge 
their  orifices  by  abrasion  of  the  older,  upper 
parts  of  the  shell  wall.  The  secondary  filling  and 


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20 


FIGURES  12-21.  Balanus  irradians  new  species.  Figs.  12-14.  Basal,  top,  and  side  views  of  paratype  CASG  60892;  greatest 
diameter  of  base  29  mm,  greatest  height  of  shell  15  mm  (note  fragment  of  basis  in  upper  part  of  Fig.  12).  Fig.  15.  Side  view  of 
shell,  paratype  CASG  60893;  greatest  height  13  mm.  Figs.  16-17.  Exterior  and  interior  views  of  scutum  tentatively  identified 
withfi.  irradians,  hypotype  CASG  60920;  height  10  mm.  Figs.  18-20.  Side,  top,  and  basal  view  of  shell,  holotype  CASG  60891; 
greatest  height  24  mm,  carinorostral  diameter  38  mm.  Fig.  21.  Top  view  of  eroded  shell  with  prominent  ribs,  paratype  CASG 
60894;  greatest  diameter  38.3  mm. 


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the  transverse  septa  of  the  parietal  tubes  prevent 
exposure  of  the  internal  tissues  of  the  barnacle 
when  the  upper  and  outer  shell  wall  are  re- 
moved. 

The  affinities  of  B.  irradians  with  B.  balanus 
and  B.  rostratus  are  in  keeping  with  the  conclu- 
sion that  the  fauna  of  the  basal  Wilson  Ranch 
beds  lived  in  hydroclimatic  conditions  similar  to 
those  of  modern  Puget  Sound.  Balanus  rostra- 
tus is  a  subtidal  North  Pacific  boreo-arctic 
species  ranging  south  to  northern  Japan  in  the 
west  and  to  Puget  Sound  in  the  east.  Balanus 
balanus  is  a  lower  intertidal  and  subtidal  species 
of  the  Arctic,  boreal  Atlantic,  and  boreal  Pacific, 
with  a  distribution  similar  to  that  of  B.  rostratus 
in  the  North  Pacific.  The  fossil  record  of  B.  ros- 
tratus is  limited  to  the  late  Pleistocene  on  the 
Pacific  coast  of  North  America  where  it  is  re- 
corded as  far  south  as  central  California  (Zullo 
1969b),  but  extends  back  to  the  Pliocene  in  Ja- 
pan (Yamaguchi  1977).  Balanus  balanus  is 
found  in  Pleistocene  deposits  of  both  the  North 
Atlantic  and  North  Pacific  basins,  but  has  not 
been  recorded  from  deposits  south  of  southern 
Oregon  (Zullo  1969b).  Miocene  and  Pliocene 
European  records  of  B.  balanus  are  spurious 
(Menesini  1968,  in  part),  but  the  species  does 
occur  in  glacio-marine  sediments  of  the  Miocene 
part  of  the  Yakataga  Formation  in  southeastern 
Alaska,  and  is  reported  from  the  Miocene  of  Ja- 
pan by  Yamaguchi  (1971). 

The  morphology  of  B.  irradians,  particularly 
those  features  peculiar  to  this  species,  suggests 
that  B.  irradians  is  a  derivative  of  either  B.  bal- 
anus or  B.  rostratus.  The  modifications  seen  in 
the  new  species  reflect  adaptations  to  life  under 
conditions  of  pronounced  wave  shock  and  abra- 
sion, perhaps  in  the  intertidal  zone  in  relatively 
open  coastal  conditions. 

Balanus  sp.  cf.  B.  irradians 

(Figures  16-17) 

A  single,  nearly  intact  scutum  from  CASG  lo- 
cality 54135  is  tentatively  referred  to  B.  irradi- 
ans. It  is  thin,  about  as  broad  as  high,  and  slight- 
ly concave  externally  between  base  and  apex. 
The  exterior  bears  closely  spaced,  semi-erect 
growth  ridges  that  are  finely  crenate.  The  tergal 
margin  is  slightly  reflexed  and  the  basitergal  an- 
gle is  rounded.  The  articular  ridge  is  convex, 
reflexed  over  the  articular  furrow,  and  protrudes 
beyond  the  tergal  margin.  The  articular  furrow 


is  narrow,  shallow,  and  short.  The  articular 
ridge  is  long,  fully  two-thirds  the  length  of  the 
tergal  margin.  A  short,  low,  blunt  adductor  ridge 
is  present  along  the  margin  of  the  large  oval  ad- 
ductor muscle  pit  in  the  upper  half  of  the  scutum 
and  is  separated  from  the  articular  ridge.  The 
depressor  muscle  pit  is  a  large  triangular  area 
between  the  adductor  muscle  pit  and  the  basi- 
tergal angle.  This  pit  is  not  bordered  by  a  ridge 
on  its  occludent  side.  The  pit  for  the  depressor 
muscles  in  the  basioccludent  angle  is  large,  tri- 
angular, and  deep. 

This  scutum  bears  some  resemblance  to  those 
of  B.  balanus  and  B.  rostratus,  but  differs  from 
both  in  its  greater  breadth,  thinness,  its  short, 
blunt  adductor  ridge  that  is  well  removed  from 
the  articular  ridge,  and  in  the  greater  size  and 
depth  of  the  depressor  muscle  pit  that  is  not  bor- 
dered by  a  ridge  on  its  occludent  side.  This  scu- 
tum differs  from  others  found  at  CASG  locality 
54135  and  identified  as  Balanus  sp.  aff.  B.  nu- 
bilus  by  its  greater  width,  its  convex  rather  than 
straight  articular  ridge  that  is  much  more  re- 
flexed  and  much  longer,  its  shorter,  blunter  ad- 
ductor ridge,  and  the  lack  of  striations  or  ridges 
in  the  depressor  muscle  pit. 

Balanus  sp. 

(Figures  31-33) 

Several  whole,  high  conic  and  cylindric  shells 
cannot  be  identified  beyond  the  generic  level. 
Their  orifices  are  toothed,  the  radii  are  moder- 
ately sunken  with  oblique  summits,  and  the  pa- 
rietes  are  smooth  to  irregularly  plicate,  to  in- 
conspicuously ribbed.  These  shells  may  represent 
either  B.  sp.  aff.  B.  nubilus  or  Notomegabala- 
nus(1)  insperatus,  but  their  preservation  is  not 
conducive  to  the  identification  of  specific  char- 
acters. 

Subfamily  MEGABALANINAE  Newman,  1980 

Notomegabalanus(?)  insperatus  new  species 
(Figures  22-30) 

DIAGNOSIS. — Shell  of  six  compartmental 
plates  with  broad,  tubiferous  radii;  summits  of 
radii  oblique;  septa  of  radial  sutural  edge  bearing 
denticulae  on  lower  sides  only;  exterior  of  pa- 
rietes  of  larger  compartmental  plates  with  three 
to  five  prominent  ribs,  and  ornamented  by  fine, 
closely  spaced  radial  striae;  parietal  tubes  nu- 
merous, rectangular,  without  transverse  septa, 


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28 


FIGURES  22-30.  Notomegabalanus(!)  insperatus  new  species.  Figs.  22-24.  Interior,  exterior,  and  right  radial  sutural  edge 
views  of  rostrum,  holotype  CASG  60914;  greatest  height  of  plate  36.4  mm.  Figs.  25-26.  Exterior  and  interior  views  of  lateral 
plate,  paratype  CASG  60915;  greatest  height  of  plate  20.5  mm.  Figs.  27-28.  Radial  sutural  edge  and  exterior  views  of  lateral 
plate,  paratype  CASG  60916;  greatest  height  of  plate  28.5  mm.  Figs.  29-30.  Side  view  and  enlargement  of  radial  sutural  edge 
of  lateral  plate,  paratype  CASG  60917;  greatest  height  of  plate  18  mm. 


ZULLO  &  GURUSWAMI-NAIDU:  NORTHERN  CALIFORNIA  MIOCENE  BALANIDS 


533 


FIGURES  31-33.    Balanus  sp.,  side  views  of  three  shells,  hypotypes  CASG  6091 1  through  60913,  respectively;  greatest  height 
of  shells:  (Fig.  31)  42  mm,  (Fig.  32)  20.5  mm,  (Fig.  33)  29  mm. 


but  secondarily  filled  in  upper  half;  basis  un- 
known, but  presumed  calcareous;  opercular 
plates  unknown;  distinguished  from  other 
species  of  Notomegabalanus  Newman,  1980, 
and  Austromegabalanus  Newman,  1980,  by  the 
prominent  external  ribbing  of  the  shell  wall. 

DESCRIPTION. — Reconstructed  shell  high  con- 
ic with  broad,  tubiferous  radii  and  moderately 
toothed  orifice;  summits  of  radii  oblique;  radial 
sutural  edges  septate,  with  septa  bearing  dentic- 
ulae  on  lower  sides  only;  alae  moderately  broad 
with  oblique  summits,  confined  to  area  of 
sheath;  length  of  sheath  less  than  one-half  height 
of  compartmental  plates;  lower  margin  of  sheath 
free-standing,  with  shallow  cavity  between  basal 
margin  of  sheath  and  interior  of  shell  wall;  ex- 
terior of  parietes  with  from  three  to  five  promi- 
nent ribs  on  the  larger  compartmental  plates, 
and  ornamented  by  fine,  closely  spaced  radial 
striae  crossed  by  weak,  widely  spaced  growth 
lines;  interior  of  shell  wall  ribbed  between  base 
and  sheath,  internal  ribs  low,  moderately  devel- 
oped, each  corresponding  to  a  parietal  septum; 
parietal  tubes  numerous,  without  transverse 
septa,  but  secondarily  filled  in  upper  half;  pari- 
etal septa  thin,  basally  denticulate;  outer  lamina 
with  one  to  four  thin,  half  or  quarter  septa;  basis 
unknown  but  presumed  calcareous  from  basal 
denticulation  of  parietal  septa;  opercular  plates 
unknown. 


MATERIAL  EXAMINED. — Twenty-four  disarticulated  com- 
partmental plates  and  one  possibly  complete  shell  from  CASG 
locality  54135. 

TYPE  MATERIAL. — Holotype,  CASG  no.  60914;  paratypes, 
CASG  nos.  60915  through  60918,  and  paratype  lot  CASG  no. 
60919  in  the  California  Academy  of  Sciences  paleontological 
type  collection. 

ETYMOLOGY. — The  specific  name  is  Latin  for 
unhoped  for  or  unexpected. 

DISCUSSION. — Recently,  Newman  (1980)  sub- 
divided the  genus  Megabalanus  Hoek,  1913, 
into  three  genera.  Megabalanus  s.s.  includes 
most  of  the  Megabalanus  tintinnabulum  (Lin- 
naeus, 1758)  complex  together  with  related 
species  of  modern  tropical  and  warm  temperate 
seas.  The  two  new  genera,  Austromegabalanus 
and  Notomegabalanus,  include  fossil  and  extant 
species  restricted  to  austral  cool  temperate  and 
subpolar  waters.  The  only  obvious  shell  char- 
acter separating  Megabalanus  s.s.  from  the  two 
austral  genera  is  that  the  septa  of  the  radial  su- 
tural edge  of  Megabalanus  are  denticulate  on 
both  their  lower  and  upper  sides,  whereas  the 
septa  of  the  austral  genera  are  denticulate  only 
on  their  lower  sides.  The  two  austral  genera  are 
separated  on  characters  of  their  terga;  that  of 
Austromegabalanus  has  a  beak  and  a  closed 
spur  furrow,  whereas  that  of  Notomegabalanus 
is  not  beaked  and  has  an  open  spur  furrow. 
There  are  features  of  the  mouth  parts  as  well 
that  can  be  used  to  distinguish  true  Megabala- 


534 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42,  No.  21 


nus  from  the  austral  genera,  but  when  dealing 
with  the  generic  assignment  of  fossil  species, 
only  shell  and  opercular  plate  characteristics  are 
available. 

The  revision  of  Megabalanus  proffered  by 
Newman  is  sound,  both  from  a  systematic  and 
biogeographic  point  of  view.  However,  the  Neo- 
gene  fossil  megabalanid  record  from  California 
presents  a  problem.  The  species  described  here 
and  "Megabalanus"  wilsoni  (Zullo,  1969a)  from 
the  Pliocene  San  Diego  Formation  have  radial 
sutural  dentitions  characteristic  of  Newman's 
austral  genera.  Furthermore,  the  tergum  of  "M. 
wilsoni  is  typical  of  Notomegabalanus,  although 
the  spur  furrow  is  closed  in  adult  specimens,  and 
its  resemblance  to  that  of  N.  algicola  (Pilsbry, 
1916)  from  South  Africa  was  noted  previously 
(Zullo  1969a). 

The  Notomegabalanus  species  closest  geo- 
graphically to  the  California  coast  is  N.  concin- 
nus  (Darwin,  1854)  from  the  Peruvian  province. 
This  species  was  not  included  by  Newman,  but 
it  has  the  sutural  dentition,  tergum,  lab  rum,  and 
protuberant  lower  margin  of  the  first  maxilla  that 
characterize  Notomegabalanus.  Are  these  Cal- 
ifornia Neogene  species  true  Notomegabalanus 
derived  from  some  austral,  perhaps  South 
American  ancestor?  Does  the  Austromegabal- 
anus-Notomegabalanus  complex  represent  an 
ancestral,  previously  worldwide  stock  from 
which  tropical  and  warm  temperate  Megabala- 
nus s.s.  evolved?  Or  do  the  California  species 
represent  a  separate  group  exhibiting  conver- 
gence of  characters  with  the  austral  genera?  At 
present  it  is  possible  only  to  consider  the  Cali- 
fornia species  on  the  basis  of  their  morphologies 
and  to  assign  them  to  Notomegabalanus  on  the 
characters  delimiting  that  taxon.  The  association 
of  N.  (?)  insperatus  with  a  cool  temperate  fauna 
does,  however,  suggest  affinities  with  the  austral 
megabalanid  genera. 

Notomegabalanus^)  insperatus  differs  from 
all  other  megabalanid  species  in  bearing  distinct, 
prominent,  external  ribs.  Megabalanus  validus 
(Darwin,  1854)  and  M.  zebra  (Darwin,  1854) 
have  ribbed  shells,  but  their  ribs  are  low,  round- 
ed, and  usually  confluent,  rather  than  erect  and 
separate.  In  addition,  both  of  these  species  have 
the  sutural  dentition  typical  of  Megabalanus  s.s. 
This  new  species  is  questionably  assigned  to 
Notomegabalanus  rather  than  to  Austromega- 
balanus  solely  on  the  basis  of  the  presence  of 


another  Notomegabalanus,  N.  wilsoni,  in  the 
California  Neogene. 

ACKNOWLEDGMENTS 

We  thank  Peter  Rodda,  California  Academy 
of  Sciences,  for  information  concerning  the  geo- 
logic setting  and  stratigraphy  of  the  Wilson 
Ranch  beds  and  for  his  review  of  the  manu- 
script; and  Barry  Roth,  California  Academy  of 
Sciences,  for  his  assessment  of  the  age  and  de- 
positional  environment  of  the  Wilson  Ranch 
molluscan  fauna  and  for  his  review  of  the  manu- 
script. 

LITERATURE  CITED 

ADDICOTT,  W.  O.  1976.  Neogene  molluscan  stages  of  Oregon 

and  Washington.  Soc.  Econ.  Paleont.  and  Mineral.,  Pacific 

Sec.,  Neogene  Sympos.,  pp.  95-115. 
BARTOW,  J.  A.,  A.  SARNA-WOJCICKI,  W.  O.  ADDICOTT,  AND 

K.  R.  LAJOIE.  1973.  Correlation  of  marine  and  continental 

deposits  in  northern  California  tephrochronology.  Am.  As- 

soc.  Pet.  Geol.  Bull.  57(4):769. 

BERGGREN,  W.  A.   1972.  A  Cenozoic  time-scale — some  im- 
plications for  regional  geology  and  paleobiogeography.  Le- 

thaia  5: 195-215. 
.  1978.  Marine  micropaleontology  an  introduction.  Pp. 

1-17  in  Haq,  B.  U.,  and  A.  Boersma,  editors,  Introduction 

to  marine  micropaleontology.  Elsevier,  New  York. 
DARWIN,  C.  1854.  A  monograph  on  the  sub-class  Cirripedia, 

The  Balanidae,  The  Verrucidae.  Ray  Society,  London.  684 

P- 
DICKERSON,  R.  E.  1922.  Tertiary  and  Quaternary  history  of 

the  Petaluma,  Point  Reyes  and  Santa  Rosa  quadrangles. 

Proc.  Calif.  Acad.  Sci.,  ser.  4,  11(19):527-601. 
HIGGINS,  C.  G.   1960.  Ohlson  Ranch  Formation,  Pliocene, 

northwestern  Sonoma  County,  California.  Univ.  Calif. 

Publ.  Geol.  Sci.  36(3):  199-232. 
HOEK,  P.  P.  C.  1883.  Report  on  the  Cirripedia  collected  by 

H.M.S.  Challenger  during  the  years  1873-1876.  Rep.  Sci. 

Results  Voy.  H.M.S.  Challenger,  Zool.  8(25):  1-169. 
.   1913.  The  Cirripedia  of  the  Siboga -Expedition.  B. 

Cirripedia  Sessilia.  Pp.  129-275  in  S&oga-Expeditie  31b. 

E.  J.  Brill,  Leyden. 
LINNAEUS,  C.    1758.   Systema  Naturae.  10th  Ed.  Holmiae. 

428  p. 
MENESINI,  E.  1968.  Balanus  curvirostratus  nuova  specie  del 

Pliocene  della  Toscana.  Atti  Soc.  Toscana  Sci.  Nat.  Mem. 

P-V,  ser.  A,  75(2):617-632. 
NEWMAN,  W.  A.  1980.  On  the  biogeography  of  balanomorph 

barnacles  of  the  southern  ocean  including  new  balanid  taxa; 

a  subfamily,  two  genera  and  three  species.  Proc.  Internatl. 

Sympos.  Mar.  Biogeogr.  and  Evol.  Southern  Hemisphere, 

New  Zealand  DSIR  Inform.  Ser.  137,  1:279-306. 
OSMONT,  V.  C.    1905.   A  geological  section  of  the  Coast 

Ranges  north  of  the  Bay  of  San  Francisco.  Univ.  Calif. 

Publ.  Bull.  Dep.  Geol.  4(3):39-S7. 
PILSBRY,  H.  A.  1916.  The  sessile  barnacles  (Cirripedia)  in  the 

collections  of  the  U.S.  National  Museum;  including  a  mono- 
graph of  the  American  species.  U.S.  Natl.  Mus.  Bull. 

93:1-366. 


ZULLO  &  GURUSWAMI-NAIDU:  NORTHERN  CALIFORNIA  MIOCENE  BALANIDS  535 

Ross,  A.  1969.  Studies  on  the  Tetraclitidae  (Cirripedia:  Tho-  ately  north  of  the  San  Francisco  Bay  region,  California. 

racica):  revision  of  Tetraclita.  Trans.  San  Diego  Soc.  Nat.  Geol.  Soc.  Am.  Mem.  35:1-242. 

Hist.  15(15):237-251.  YAMAGUCHI,  T.  1971.  Fossil  barnacles  from  the  Pleistocene 

ROTH,  B.,  AND  R.  GURUSWAMI-NAIDU.  1978.  A  new  Mio-  Miyata  Formation.   Sci.   Rep.   Yokosuka  City   Mus. 

cene  species  of  Nuttallia  (Mollusca:  Bivalvia)  from  the  Sa-  18:122-130. 

Unas  Valley,  California.  J.  Paleont.  52(l):61-66.  .  1977.  Taxonomic  studies  on  some  fossil  and  recent 

SARNA-WOJCICKI,  A.  M.  1976.  Correlation  of  late  Cenozoic  Japanese  Balanoidea.  Trans.  Proc.  Palaeontol.  Soc.  Jpn, 

tuffs  in  the  central  Coast  Ranges  of  California  by  means  of  new  ser.  107-108:135-201. 

trace-  and  minor-element  chemistry.  U.S.  Geol.  Surv.  Prof.  ZULLO,  V.  A.   1969a.  Thoracic  Cirripedia  of  the  San  Diego 

Pap.  972: 1-30.  Formation,  San  Diego  County,  California.  Nat.  Hist.  Mus. 

SCHUMACHER,  C.  F.  1817.  Essai  d'un  nouveau  systeme  des  Los  Angeles  Cty.  Contrib.  Sci.  159:1-25. 

havitations  des  vers  testaces.  Copenhagen.  287  p.  .  1969b.  A  late  Pleistocene  marine  invertebrate  fauna 

TRAVIS,  R.  B.  1952.  Geology  of  the  Sebastopol  quadrangle,  from  Bandon,  Oregon.  Proc.  Calif.  Acad.  Sci.,  ser.  4, 

California.  Calif.  Div.  Mines  Bull.  162:1-33.  36(1):347-361. 

VAN  EYSINGA,  F.  W.  B.  1975.  Geological  time  table,  3rd  Ed.       .  1979.  Thoracican  Cirripedia  of  the  Pancho  Rico  For- 

Elsevier,  Amsterdam.  mation,  Salinas  Valley,  Monterey  County,  California.  Nat. 

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CALIFORNIA  ACADEMY  OF  SCIENCES 

Golden  Gate  Park 
San  Francisco,  California  94118 


INDEX  TO  VOLUME  42 


(Compiled  by  Lillian  J.  Dempster) 
New  names  in  boldface  type. 


kbraliopsis  425,  428 
Acacia  76,  82 
Kcanthurus  triostegus  65 
{chillea 

Millefolium  20,  24,  31,  33,  45 

sp.  20,  24 
\cmaea  324-325,  328 

striata  337 
Acmaeidae  323-339 
(cmonotus  190 

chilensis  190 
(cyphoderes  70,  72-73,  80 

acutipennis  73,  78-79 

amoena  69,  73-76 

aurulenta  72 

cribricollis  73-74 

delicata  73,  78 

delicatus  78 

fulgida  69,  73,  76-77 

parva  69,"  73,  77,  79-80 

prolixa  69,  73,  78 

sexualis  73,  76-77 

suavis  73,  78,  80,  82 

velutinus  72 

vespiventris  72 

yucateca  73,  79 
tg/a/a  fidelis  var.  minor  398 
.{glaophenis  latirostris  427 
Agonidae  429 
-Agonina  458 
Ifl/wm  20 

; :  l/oja  sapidissima  410 
\\mblyarrhena  305-306 
;\<iaphalis  sp.  45 
\ndira  inermis  72 
Unillina  455,  458-459,494 
|tn///u£  integripennis  485,  487 
\noplopoma  fimbria  423 
%timora  341-348 

oust  rails  341-343,  348 

meat//  341-342,  348 

microlepis  341-344,  347 

rhino  341-343,  347 

rostrata  341-344,  346-348 

spp.  342 
to/ea  acus  61 
tyhonus  4 

pyriformis  50 
ztyistus  287 
•\  \Ploactinidae  287 
Wloactis  288 


Apristurus  brunneus  414,  419 
Apterichtus  flavicaudus  57-58,  63,  66 
Araneae  318 
Arceuthobium 

cyanocarpum  3,  20,  24 

sp.  24 

Archaeogastropoda  323 
Archoleptoneta  schusteri  318 
Archytas  10 

Arctocephalus  townsendi  411 
Arctophyto  11 
Arctophyto-Milada  11 
Arctostaphylos  384 

sp.  24 

Ardistomina  458 
Anemia  214 
Aruncus  sp.  23 
Ascothoracica  443 
Ascothoracida  443-444 
Ascothorax  444 
Aster  sp.  24,  45 
Asteroidea  443 
Ateloglossa  9,  1 1-12 
Austromegabalanus  533-534 
Austromegabalanus-Notomegabalanus  complex  534 


Baccharis  3 

glutinosa  20,  49 

pilularis  20,  33 

sp.  20,  24,  33,  49 
Balanidae  525,  527 
Balaninae  527 
Balanoidea  527 
Balanus  529 

ftcr/an«j  525,  529,  531 

balanus  complex  525,  529 

irradians  525,  529-531 

nubilus  525,  527,  529 

proxinubilus  525,  529 

rostratus  529,  531 

sp.  531,  533 

sp.  aff.  B.  nubilus  525,  527-529,  531 

sp.  cf.  B.  irradians  531 
Baltimore  80 
Banksula  315-322 

californica  315-317,  322 

calif  or  nica  group  316-319 

elliotti  315-317,  319-322 

galilei  316-317,  322 

grahami  315-317,  320-322 

grubbsi  315-317,  319-320,  322 


[537] 


538 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42 


martini  317 

martinorum  315-319,  322 
melones  315-317,  320-322 
melones  group  316 
rudolphi  315-318,  322 
tuolumne  316-317,  332 
IBelone  caudimaculata  301 
Bembidiina  455,  458 
Bembidiini  455-456,  458-459 
Bembidion  455-456,  458-460,  462-464 

aeger  455,  460,  462-466,  470,  473,  475-476,  478, 

491-492 

affine  group  460,  482 
aratum  460-464,  478-480,  491,  493 
armuelles  455^60,  462-465,  467,  481-482,  491,  493 
barrens!*  455,  460-464,  467,  470,  482-483,  491, 

493 

championi  460-464,  467,  483-484,  491,  493 
championi  group  460,  483 

chiriqui  455,  460,  462-466,  470,  476,  478,  491-492 
cognatum  group  460,  484 
cortes  455,  460,  463^t65,  467,  470,  478,  480-481, 

491,  493 

cyclodes  460-464,  483,  491,  493 
(Cyclolopha)  483 
definitum  479 
diabola  455,  460,  462-466,  470,  473-475,  478,  491- 

492 

dorsale  group  460,  480 
edwardsi  455,  460, 462-466, 470,  473,  475-476, 491- 

492 

(Eupetedromus)  479-480 

franiae  455,  460-464,  467,  470,  477,  479,  491,  493 
franiae  group  460,  477 
(Furcacampa)  482 
(Hydromicrus)  465 
incrematum  group  460,  479 
ixtatan  455,  460-464,  467,  470,  478,  484-485,  491, 

493 
lavernae  455,  460,  462-466,  472-473,  475-476,  491- 

492 

mexicanum  460,  462-466,  469-471,  478,  491-492 
nahuala  455,  460-464,  467,  470,  477-478,  491,  493 
nahuala  group  460,  476-477 
(Notaphus)  479-480 
(Peryphus)  466,  468,  477 
purulha  455,  460,  462^67,  470,  491-492 
quetzal  455,  460,  462-466,  470,  473-474,  478,  491- 

492 

rogersi  460-464,  466-468,  470,  478,  491^92 
rogersi  group  460,  467 
sanctaemarthae  467 

satellites  460,  462-466,  470-472,  478,  491-492 
semistriatum  group  460,  465-466 
sparsum  460-464,  467,  478,  481^82,  491,  493 
subapterum  467 
transparens  group  484 


vernale  457,  460,  462-466,  469-472,  478,  491-492 

vernale  group  460,  467-469,  477 

versicolor  group  482,  484 

vinnulum  479 

vulcanium  460,  462-466,  470,  474-476,  478,  491- 

492 

Bembidium  459 
affine  482 
aratus  479 
aztecanum  485 
championi  483 
cyclodes  483 
mexicanum  469 
rogersi  467 
sallaei  469 
satellites  471 
scintillans  479 
semistriatum  465 
sparsum  481 
sphaeroderum  483 
tetracolum  468 
vernale  469 
Bigelovia  sp.  24 
Bixa  70 
Blenniidae  429 
Brachysomophis  64 

henshawi  57,  59,  64,  66 

sauropsis  57,  59,  64,  66 
Bromiades  70,  72 

brachyptera  72 

meridionalis  72 
Bryx  182,  184,  192-193,  197-199,  220,  222 

arctus  181,  184,  193-194,  1%,  220-221,  223 

balli  193,  220 

banned  193,  220 

(Bryx)  192-193,  197,  220,  222 

(Bryx)  clarionensis  198 

(Bryx)  veleronis  197-198 

clarionensis  181,  184,  193,  197-199 

coccineus  181,  184,  193-194,  196,  220-221 

darrosanus  193,  220 

dunckeri  184,  197 

herald!  181,  192-194,  1%,  220 

hildebrandi  193,  1% 

randalli  197 

(Simocampus)  192-193,  195-196,  220,  222 

(Simocampus)  arctus  193,  195-1% 

(Simocampus)  balli  193 

(Simocampus)  banned  193 

(Simocampus)  coccineus  193,  196-197 

(Simocampus)  darrosanus  193 

(Simocampus)  herald!  192-195 

(Simocampus)  hildebrandi  193 

veleronis  181,  184,  192-193,  197-199 
Buddleia  80,  82 

wrightii  80 
Bulimulidae  399 
Byrsonima  70 


.INDEX 


539 


"acalia  decomposita  24,  26 
"aecula 

platyrhyncha  63 
(Sphagebranchus)  flavicauda  63 
(Sphagebranchus)  platyrhyncha  63 
'alathea  483 
*allechelys 

galapagensis  63,  66 
luteus  57-58,  63,  66 
>llionymidae  349 

•allionymus  349-353,  357,  359-361,  363-364,  366,  369, 
371,  373,  375 

africanus  349-354,  362,  364-366 
altipinnis  349-352,  355,  373-375 
bentuviai  349-350,  352,  354,  366-367 

calauropomus  359 

(Callionymus)  349-350,  364 

(Calliurichthys)  349 

carebares  349-351,  353-354,  356-357,  376 

curvicornis  370,  373 

formosanus  349-352,  355,  369-370 

guentheri  349-353,  355,  361,  370-373 

huguenini  373,  375 

indicus  349 

kaianus  349-353,  355,  357-361,  363-364,  366,  369- 
370,  372-373,  375 

kaianus  -group  349-377 

kaianus  moretonensis  349,  359 

kotthausi  349-352,  354,  362-364 

lyra  349 

maculatus  349 

moretonensis  349-353,  355,  358-360,  369,  376 

ochiaii  349-352,  355,  366-369,  373,  375 

regani  349-351,  353-354,  360-363,  366,  372-373 

sokonumeri  349-352,  355,  375 

sp.  375-376 

(Spinicapitichthys)  349 

whiteheadi  349-350,  352,  354,  359-362,  372 
llorhinus  412,  414-415,  417-418,  428 

ur sinus  411,  423 
Jnaenidae  399 
=-prifoliaceae  45 
Jabidae  87,  97,  435,  455 
\rabus 

dentellus  479 

gyllenhali  87,  111 

litorale  468 

lividus  113 
|  nivalis  87,  111 

rufescens  111 
I  rufescens  gyllenhali  111 

varius  480 
'eproctus  421 
*aria  72,  80 
nitida  77 

othus  46 
omericanus  45 


fendleri  24 
sp.  24,  25 

Cephaloscyllium  418 
Cerambycidae  69,  84 
Cetonurus  135,  142-143,  147 
Chaenopleura  309-312 
Characidae  499 
Charis  71 

euphrasy ne  71 
Charisia  71 

nigerrima  71 
Chauliodus  eximius  218 
Chilara  418,  421 

taylori  409 
Chiroteuthidae  427 
Chiroteuthis  426,  428 

sp.  421 
Choeroichthys  222 

brachysoma  184 

Chrysanthemum  leucanthemum  45 
Chrysothamnus 
greeni  24 
viscidiflorus  20,  33 
Cicindela  quadrimaculata  459 
Cirrhimuraena 
macgregori  64 
playfairii  57,  59,  64,  66 
Cirripedia  443,  525,  527 
Cirsium  sp.  27,  45,  51 
Clevelandia  ios  219 
Clinidae  429 
Clupea 

harengus  423 
hare ng us  pallasii  416 
Clupeidae  299 
Coccoloba  72 
Coecilophis  62 
Coelocephalus  144 

acipenserinus  144 
Coelorinchus  135-136,  144,  156 
Coleoptera  69,  87,  122,  435 
Collembola  321 

Collisella  324-326,  328,  336-337 
acutapex  328 
atrata  326 
discors  326 
mitella  326,  328,  337 
pediculus  326,  335-336 
pelta  332 
triangularis  328 
mrver/  328 
Cololabis  saira  423 

Compositae  20,  23-26,  33,  41-42,  45,  49,  51 
Convolvulaceae  46 
Cordia  70,  72 
Coryphaenoides  135 
garmani  151-152 
misakius  153 


540 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol  42 


Corythoichthys  222 
flavofasciatus  184 

intestinalis  223 

sp.  184 

"Corythoichthys"  222 
Cosmocampus  192,  222 

brachycephalus  184,  223 
Cottidae  429 
Crago  sp.  414,  417-418 
Cremanium  303,  311 
Cm  ton  74-75,  82 

sp.  49 

Cruciferae  31,  49 
Crustacea  443 

Cymatogaster  aggregate  416 
Cypriniformes  497,  500 
Cystophora  429 

Dalea 

albiflora  49 

sp.  24 

Daucus  carota  45 
Delphinidae  411 
Delphinus  412^13,  416-418,  425,  428 

delphis  411  ,425 
Dendrogaster  443-444,  447-448,  450,  453 

arbusculus  443-448,  450,  452-453 

cf.  arbusculus  448,  450 

arctica  450 

astropectinis  448 

beringensis  448 

dogieli  448 

fisheri  443-444,  448-450,  452 

iwanowi  448 

ludwigi  448,  450,  453 

murmanensis  447 

okadai  448,  453 

punctata  443-444,  450-453 

repertus  447 

rimskykorsakowi  453 

tasmaniensis  448 
Dendrogastridae  444 
Dentirostrum  222 

janssi  184 
Dermatostethus  198 

punctipinnis  185,  198 
Dexia  16 

albifrons  1,  42,  44 

canescens  38 

cerata  1,  42,  44 

halone  39 

harpasa  24,  37,  41 

prexaspes  40 

rufipennis  36,  42,  44 
Dexiosoma  fumipennis  1,  48 
8,  11-12,42 

robust  a  1,  41-42 


Diplogrammus  350,  364 

africanus  350,  364,  366 

J/W/JCM.S  350,  363-364 
Diptera  1,  14 
Dolichocodia  11 
Doryichthys  californiensis  188 
Doryrhamphus  182,  184,  187-189,  222 

californiensis  188 

ejccww*  187,  190 

melanopleura  181,  184,  187-190,  221 

melanopleura  pleurotaenia  189 

negrosensis  187 

paulus  181,  188-190,  221 
Dosidicus  gigas  426 
Dunckerocampus  222 

dactylophorus  184 


Echidna  uniformis  61 
Echinomacrurus  135,  142-143 

mo//w  142,  144 

occidentalis  135,  138,  142-144 
Efferia  aestuans  41 
£/o/?.s  295 

Embiotoca  jacksoni  417 
Embiotocidae  416-417 


farinosa  24 

sp.  24 
Engraulis  409,  421 

mordax  409 
Enhydra  lutris  411 
Enneanectes  sexmaculatus  213 
Enoploteuthidae  427 
Entelurus  218 
Eogastrophinae  216 
Eopsetta  421 
Epimelitta  70-72 

aureopilis  71-72 

meliponica  71 

nigerrima  71 
Eptatretus  418 

J/OM/H  415,  418 
Ericaceae  24 
Eriogonum  31 

elatum  33 

fasciculatum  36 

nudum  20 

sp.  20,  24,  31 
Estheria  16 

/ttfo/b  16-17,  36-37 
Eumetopias  410,  412,  414-415,  417-421,  428-429 

jubatus  409,  411,419-421 
Eupatorium  41 

coelestinum  42 

perfoliatum  45 
Euphorbiaceae  49 


HDEX 


541 


:abaceae  24,  42,  45,  49 
•arlapiscus  185 
grsteronia  70,  72 
ragaria  107 

iadidae  176 

jadiformes  341 

flaws'  macrocephalus  417 
merlangus  423 
morhua  423 

arrinae  499 

istromyzon  497-515,  523 
borneensis  497-504,  509-512,  514 
contract™  497-502,  504-505,  512,  514-515 
ctenocephalus  497-498,  501-502,  505-507,  514 
fasciatus  498-502,  507-510,  514 
lepidogaster  497,  501-502,  508-510,  514 
megalepis  497,  499,  501-502,  509-511,  514 

[   monticola  497-499,  501-504,  511-512,  514-515 

f    punctulatus  501-502,  512-515 

;    ridens  497,  501-502,  504,  512-515 

istromyzontinae  497-501,  516 

btrophori  184,  218,  222-223 

istropoda  379 

\;ocharidius  455-456,  458,  485,  487 
gimlii  455,  485,  487-488,  491,  493 
integripennis  485,  487-488,  491,  493-494 
phineus  455,  485-487,  489-491,  493^94 
romeoi  455,  485,  487-488,  491-492 
tagliantii  455,  485,  487-488,  491,  493-494 
raea  canescens  24 

\miopsis  497-500,  515-519,  521-523 
denudata  497,  516-518 
gossei  497,  516,  518-519 
hanitschi  515-516,  518-523 
multiradiata  497,  515-516,  518,  520-523 

}hicephala  427 

Zptocephalus  418,  421 

\  zachirus  417 

faatidae  426-428 
wtus  425-426 

I  sp.  425,  428  ' 
uypium  sp.  49 

Wampus  412-413,  417-418,  427 

?  griseus  411,  426-427 

*;  stearnsii  426 

fierrezia 

r  sarothrae  25 

I  sp.  24 

ropleurodus  quoyi  282 

mutichthys  235 

foporphyrus 

f  rostratus  314-343,  347 

I  viola  341-343,  347 

Wopappus  sp.  20 

mnthus  sp.  24 


Heliopsis 

parvifolia  24-26 

sp.  25 
Helix  fidelis  379,  381 

hillebrandi  382 
Hellenium  hoopesii  27 
Helminthoglyptidae  400,  402 
Helobia  castanipes  87,  111-112,  115 
Hemilepidotus  hemilepidotus  423 
Hemithylacus  Peter  si  190 
Heraldia  222-223 

nocturna  184 
Hesperodinera  8,  10-12 
Hexagrammos 

decagrammus  409,  417 

lagocephalus  416 
Hippasteria  californica  443—444 
Hippocampus  182-187,  221-223 

breviceps  185 

ecuadorensis  186 

gracilis  186 

heptagonus  185 

hildebrandi  181,  186-187 

ingens  181,  184-187,  197,  221,  224 

kuda  184 

reidi  187 

spinosissimus  185 
Hippoglossoides  elassodon  416 
Hippohystrix  185 
Hipposyngnathinae  216-217 
Hipposyngnathus  216-218,  221 

convexus  217-218 

imporcitor  181,  217-218 

neriticus  218 
Histioteuthidae  427 
His  trio  his  trio  185 
Homalopteridae  497 
Horologion  455,  459 

speokoites  459 
Hydrolagus  418 
Hymenocephalus  135-137,  140,  149 

gracilis  140 

(Hymenogadus)  140 

(Hymenogadus)  gracilis  140 

(Hymenogadus)  tenuis  140 

italicus  140 

kuronumai  140 

sp.  137,  140-141 

(Spicomacrurus)  140 

(Spicomacrurus)  kuronumai  140 

tenuis  140 
Hymenogadus  140 
Hypsoblennius  gentilis  219 
Hystrichodexia  16 

contristans  16,  25-26 

Ichthyapus  vulturis  57-58,  63,  66 


542 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42 


Ichthyocampus  222 

belcheri  184 

Icichthys  lockingtoni  425 
Ilypnus  gilberti  219 
Ipomoea  46 

Jatropha  74,  80,  82 

curcas  75 
Jenkinsiella  macgregori  64 

Kogia  412-413,  417-418,  427 

breviceps  427 

simus  411,  427 
Kumba  135,  147 

Labeoinae  499 
Labiatae  49 
Laemonema  425 

Lagenorhynchus  409,  412-413,  416-418,  425-426,  428, 
430 

obliquidens  409,  411,  425-426 
Lampetra  tridentata  418 
Leiuranus  semicinctus  59,  66 
Lepidium  thurberi  49 
Lepidoglanis  500 

monticola  500,  511 
Lepidospartum  squamatum  20 
Leptasterias  groenlandica  450 
Leptenchelys  labialis  62 
Leptocottus  armatus  416 
Leptonotus  181-182,  184,  190,  192,  221-222 

blaenvillianus  190 

blainvilleanus  181,  184,  188,  190-192,  221 

Blainvillei  190 

blainvillianus  190 

elevatus  192 

norae  192 

semistriatus  192 
Leucichthys  424 
Liliaceae  20 
Liocranoides  sp.  318 
Lionurus  156 

cetonuropsis  147 

liolepis  157 

(Lionurus)  barbiger  157 

(Lionurus)  fragilis  147 

(Lionurus)  liolepis  157 

(Nezumia)  convergens  171 

(Nezumia)  cuspidatus  171 

(Nezumia)  latirostratus  169 

(Nezumia)  loricatus  174 

(Nezumia)  orbitalis  167 

(Nezumia)  pudens  163 

(Nezumia)  stelgidolepis  160 

(Nezumia)  trichiurus  171 

nigromaculatus  153 

stelgidolepis  160 

ventralis  167 


Lippia  wrightii  24 

Lo//go  409,  413,  418,  424-425,  430 

opalescens  421-422,  424,  428-429 
Lonicera  sp.  45 
Lophiiformes  229 
Lophius 

histrio  243 

(Malthe)  cubifrons  258-260 

nostratus  274 

radiatus  258-260 

rostratus  251-253,  260,  274 

vespertilio  234,  247,  251-252 
Loranthaceae  24 
Lott/a  323-326,  333-334,  337-338 

gigantea  325-326,  332,  335-336 

mesoleuca  323,  325,  336-338 

mimica  323,  325-326,  332-338 

mimica-smithi  335,  337 

smith!  323,  326,  333,  334-338 

stipulata  336 

Lycodopsis  pacifica  409,  417 
Lymnastis  459 
Lyopsetta  418,  421 

Macrocystis  206 

angustifolia  207 

pyrifera  207,  220 
Macrodactylus  4 

subspinosus  1,  4,  42 
Macrometopa  49 

calogaster  48 

Macrouridae  135-136,  177,  427 
Macrourinae  135 
Macrourus  135 

/aevtt  149 

lucifer  152 

nigromaculatus  152-153 

nigromarginatus  152 

serratus  156 
Macruroplus 

ori  152 

potronus  153,  155,  176 

pudens  163 

pulchellus  159 
IMacruroplus  156 

nigromaculatus  153 
Macrurus  140,  149 

barbiger  157-159 

convergens  171,  173 

cuspidatus  171,  173 

fasciatus  152 

flagellicauda  147 

/rag///;  135,  147 

gracillicauda  160 

(Hymenogadus)  140 

latirostratus  168 

//otepw  157,  159 

(Lionurus)  liolepis  157 


INDEX 


543 


loricatus  174 
(Malacocephalus)  149 
(Malacocephalus)  laevis  149 
(Mystaconurus)  140 
orbitalis  167 
parvipes  147 
stelgidolepis  160 
tenuicauda  145 
trichiurus  171,  173 
\\alacocephalini  157 
Malacocephalus  135,  149,  151,  157 
hawaiiensis  149 
/flevw  137,  149-151 
nipponensis  149,  151 
sp.  149 

iallotus  villosus  423 
ialthaea  234 

angusta  243,  246-247 
cubifrons  258 
longirostris  247,  250 

247,  251,  258,  267 
243,  247 
truncata  243,  247 

234 
angusta  243 
cubifrons  258,  260 
nasMta  251,  253,  258 
notata  243 
radiata  258 
truncata  243 

vespertilio  234,  243,  247,  258,  274,  276 
vespertilio  nasuta  258 
vespertilio  radiata  258 
to/iea  234 
guacucuja  247 
nasuta  258 
notata  243 
radiata  258 
vespertilio  251 
'dthopsis  234 
alvaceae  49 
wwbra  218,  222 
pers  errata  184 
vrubium  vulgare  49 
itaeocephalus  135,  144-145 
acipenserinus  146-147 
adjustus  135,  144-145,  147 
microstomus  147 
nigrescens  146-147 
tenuicauda  138,  145-147,  177 
tt(7//cas/a  221 
'dicago  saliva  49 
^abalaninae  531 
abalanus  533-534 
tintinnabulum  533 
tintinnabulum  complex  533 
534 


wilsoni  534 

ze^ra  534 

"Megabalanus"  wilsoni  534 
Melastomataceae  303,  305 


42,  45 

sp.  24 

Melolonthinae  3 
Merluccius  409 

productus  409,  416-418 
Mesobius  135,  141 

antipodum  141-143 

feerry/  137,  141-143 
Miconia  303-314 

andreana  306 

chionophylla  310-311 

chiriquiensis  303-305 

coloradensis  303,  305-307 

concinna  303,  305,  307-309,  314 

confertiflora  303,  309-312 

g#6a  306-307 

grandidentata  303,  311-312 

longibracteata  303,  308-309,  312-314 

micrantha  307 

myrtillifolia  308 

parvifolia  311 

/•«&*?/!,$  305,  308 

superposita  314 

tabayensis  307 

w/ttii  307 
Micratopus  459 
Microdonophis  64 

erabo  65 

fowleri  64-65 

macgregori  64 

polyophthalmus  64 
Microgadus  409 

proximus  416-417 
Micrognathus  222-223 

fta/fi  192 

brevirostris  184 

crinigerus  214 
Microphis  extensus  189 
Microstomus  421 


Mioptachys  458 

Mirounga  412,  414,  417-418 

angustirostris  411,  418-419 
Mochlosoma  5-12 
Mollusca  323,  423 
Monadenia  379-383,  385,  390,  394,  396,  398-403 

antecedens  401 

chaceana  380,  394,  397,  399 

c/mrc/z/  378-380,  382-383,  385,  387-388,  395-3%, 
398 

circumcarinata  400,  402 

(Corynadenia)  379-380,  382-383,  400-403 

(Corynadenia)  hirsuta  382 


544 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol.  42 


(Corynadenia)  mormonum  385,  401 

cristulata  382 

dubiosa  401 

fidelis  378,  382-383,  387,  390,  398-401 

fidelis  leonina  399 

fidelis  minor  398-399 

fidelis  pronotis  400 

fidelis  scottiana  400 

hillebrandi  399,  402 

infumata  382 

marginicola  401 

marmarotis  382 

(Monadenia)  379-380,  382-383,  400-403 

(Monadenia)  fidelis  402 

(Monadenis)  setosa  402 

mormonum  385,  399 

mormonum  cala  398 

mormonum  group  401 

sefcwa  382-383,  399-400,  402 

(Shastelix)  379,  381-383,  395,  400-403 

(Shastelix)  churchi  382-402 

(Shasteftx-Corynadenia)  stock  401-403 

(Shastelix)  troglodytes  chaceana  397-399 

(Shastelix)  troglodytes  troglodytes  395-396 

(Shastelix)  troglodytes  wintu  396-397 

troglodytes  379-407 

troglodytes  chaceana  379-380,  389,  391-394,  397- 
399,  406 

troglodytes  wintu  379,  383,  386-389, 391-392,  394, 

396-398,  406 
Moridae  341,  425,  427 
Morone  saxatilis  410 
Moroteuthis  sp.  429 
Muraena  maculosa  63 
Muraenichthys  61,  221 

breviceps  61 

chilensis  60 

cookei  57-58,  63,  66 

devisi  61 

gymnotus  58,  66 

labialis  62 

laticaudata  63 

macropterus  57,  61 

ogilbyi  61 

owstoni  61 

puhioilo  57-61,  65-66 

schultzei  58,  66 

(Scolecenchelys)  61 
Mustelus  californicus  424 
Myceropsis  11,  15,  54 
Myctophidae  424-425 
Myliobatidae  418 
Myoxocephalus  sp.  416 
Myrichthys  63 

6/eeter/  57,  59,  64,  66 

maculosus  57,  59,  63-64,  66 

magnificus  63-64 


pantostigmius  57,  63 

stypurus  64 

tigrinus  57,  63 

xysturus  57,  63 
Myriocladus  444,  448 

arbusculus  444 
Myrophinae  57-58,  62 


87-133,  435-442 

98,  131 

quileute  87,  98-99,  122,  126,  128,  131 
arkansana  100,  112,  131 

arkansana  edwardsi  87,  100-102,  125-126,  128,  131 
arkansana  ooowah  87,  102,  126,  128,  131 
arkansana  uinta  87,  102-103,  125,  127,  131 
to/aria  87,  96,  112 
earn  87,  105,  121,  123,  132 
castanipes  114 
catenata  87,  112 
columbiana  108 
danmanni  435-440,  442 
darlingtoni  87,  104,  123,  132 
diver  sa  112-113 
elias  90,  112 

eschscholtzii  87,  90,  112,  115 
expansa  87,  112 
fragilis  131 
fragilis  fragilis  131 
/rag/fa  teewinot  87,  103,  127,  131 
frigida  116 
fusiformis  112 
gebleri  87,  105,  113,  132 
geWeri  cascadensis  87,  105-106,  123,  132 
gebleri  fragariae  87,  106-107,  123,  127-128,  132 
gebleri  gebleri  132 
gebleri  rathvoni  115 
gebleri  siskiyouensis  87,  107,  123,  132 
gouleti  87,  94-95,  121-122,  129 
gregaria  87,  113 
gyllenhali  90,  97,  111,  130 
gyllenhali  castanipes  111-112,  114-115,  130 
gyllenhali  gyllenhali  111 
gyllenhali  lassenensis  87,  96-97,  121,  135 
gyllenhali  lindrothi  87,  97-98,  121,  130 
hudsonica  87,  113 
incerta  87,  113 
ingens  87,  113 
kincaidi  107,  132 

kincaidi  ball!  87,  107-108,  121,  125,  132 
kincaidi  kincaidi  132 
lacustris  94-95,  112 
lacustris  bellorum  87,  95-%,  122,  129 
lacustris  lacustris  129 
Iituyae87,  100,  124,  131 
tivida  87,  112-113 
longula  87,  113 
mannerheimii  87,  114-115 


vJDEX 


545 


meanyi  109,  133 

meanyi  giulianii  435-439,  441-442 

meanyi  lamarckensis  87,  109,  125,  133 

meanyi  meanyi  133 

meanyi  sylvatica  87,  109,  124,  133 

melsheimeri  90 

metallica  87,  114 

moesta  87,  114 

molbis  114 

mollis  87,  114 

muehlenbergii  114 

miihlenbergii  114 

navajo  87,  104-105,  127,  132 

nivalis  97,  111,  130 

nivalis  bifaria  96 

nivalis  gaspesiana  87,  96,  122,  130 

nivalis  nivalis  112,  114,  130 

obliqua  87,  113-114,  116,  132 

obliqua  chuskae  87,  105,  123,  125,  132 

obliqua  obliqua  132 

obtusa  87,  114-115 

oregona  87,  115 

ovipennis  87,  115 

pallipes  114-115 

picicornis  113 

rathvoniSl,  115,  132 

sahlbergi  97,  100 

sahlbergii  87,  97,  115-116,  131 

sahlbergii  modoc  87,  99,  123,  131 

sahlbergii  sahlbergii  116,  131 

sahlbergii  triad  87,  99,  123,  131 

schwarzi  133 

schwarzi  beverlianna  87,  110,  124-125,  133 

schwarzi  schwarzi  133 

sonorae  435-442 

spatulata  112 

spatulata  sierrae  87,  108-109,  121,  133 

spatulata  spatulata  112,  133 

suturalis  87,  96,  113,  115 

tenuipes  87,  116 

texana  116 

rn/or/a  87,  116,  133 

trifaria  catenata  112,  133 

trifaria  trifaria  133 

rn/ar/a  utahensis  87,  90,  110-111,  121,  124,  127- 

128 

trifaria  vandykei  109 
turmaduodecima  435-440,  442 
vandykei  87,  109,  116,  133 
vandykei  vandykei  133 
vandykei  wyeast  87,  109-110,  121,  133 
violacea  87,  115-116 
wr/rfis  87,  116 
lioni  132 

z/om  oasis  87,  103-104,  121,  132 
lioni  zioni  132 
iini  87-133,  435-442 


Neogastromyzon  497,  499-501 

nieuwenhuisi  499,  501 

pauciradiatus  493,  505 
Nerophis  182,  218,  222-223 

Nezumia   135,  144-145,  151-152,  156-160,  163,  165, 
167,  169,  171,  173-174,  176 

aequalis  168 

africana  158,  163 

atlantica  158,  163 

bubonis  158,  163 

burragei  144,  158,  171 

condylura  156 

convergens  139,  156-157,  169-177 

convergens-loricata-orbitalis  complex  176 

cuspidata  171 

cyrano  171 

dara  158,  171 

ectenes  171 

gracillicauda  160 

hebetata  158,  171 

holocentrus  158,  171 

kamoharai  158 

latirostrata  138,  140,  146,  156-157,  168-177 

latirostratus  171 

w  137-139,  154-159,  162-163,  165,  177 
135,  139-140,  156-157,  171-172,  174-176 

loricata  atomos  135,  169,  174-177 

loricata  loricata  135,  174-175 

macronema  158 

nigromaculata  153 

obliquata  171 

/w  139,  156-157,  164,  167-168,  172-173,  176 
m  138,  141,  156-157,  162,  176-177 

propinqua  167,  171 

proximo  171 

pjute/is  139-140,  156-157,  160,  163-165,  177 

pulchella  139,  154-157,  159-160,  163,  165,  177 

sclerorhynchus  165,  167 

spp.  163 

stelgidolepis  139,  154-158,  160-163,  165,  173,  176- 
177 

suilla  165,  167,  173 

tomiyamai  171 

trichiura  171 

ventralis  135,  157,  164-167,  177 
Nimioglossa  10-12 
Notoacmea  323-326,  328,  330,  337-338 

fascicularis  325 

yi/osa  323,  330,  332,  337-338 

immaculata  323,  325,  329-332,  335-336,  338 

insessa  337 

persona  332 

pumila  323,  325,  327-329,  335-336,  338 

rothi  323,  325,  329-332,  335-336,  338 

rothi-immaculata  complex  332 

ubiquita  323,  325-329,  335-336,  338 


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Notomegabalanus  533-534 

algicola  534 

concinnus  534 

wilsoni  534 

Notomegabalanus(l)  insperatus  525,  531-534 
Novacampus  190 

Octopoteuthidae  426-428 
Octopus  409,  425,  429 

sp.  415,  417,  421,  428 
Odontocera  70,  80-91 
aurocincta  81 

aurocincta  arizonensis  82-83 
aurocincta  aurocincta  81-83 
aurocincta  var.  nigrocapicalis  81 
brachyptera  72 
carinicollis  82 
clara  81-83 
fuscicornis  81 
vitrea  81 
yucateca  79 

Ogcocephalidae  229-231,  234,  243 
Ogcocephalus  229-285 

corniger  229,  235-241,  245,  247-249,  254,  261,  265, 

268,  270,  272,  274-276 
cubifrons  229,  231-233,  235-240,  242,  245,  247- 

249,  252-255,  257-268,  274,  276 
danvini  229,  234-240,  243,  245,  248,  279-283 
declivirostris  229,  235-241,  245,  248,  261-262,  264- 

272,  276-277 
nasutus  229,  235-240,  242-243,  245-261,  264-265, 

268,  272,  274,  282 
nasutus  var.  cayennensis  243 
notatus  229,  235-249,  254,  265-267,  270,  272-273 
pantostictus  229,  235-240,  242,  245,  247-249,  253- 

255,  257,  260-262,  264-268,  270 
parvus  229,  235-241,  243,  245,  248,  254,  260-261, 

265,  268,  270-272,  274,  276-279 
porrectus  229,  235-240,  243,  245,  248,  279-283 
pumilus  229,  235-243,  245, 247-248, 254, 270-274, 

276 

radiatus  229,  243,  246,  252,  259,  264 
rostellum  229,  235-240,  242,  245,  248-249,  253- 

254,  256-257,  260-262,  266-269,  274 
sp.  256,  269,  274,  276 
truncatus  243 
vespertilio  229,  235-240,  242-251,  253,  255,  259, 

263-265,  270,  274,  276,  282 
Ommastrephes  424 
Ommata  70,  80,  83-84 
championella  84 
cyanea  84 

(Eclipta)  championella  84 
(Eclipta)  cyanea  84 
elegans  83 

(Ommata)  rubriventris  84 
sallaei  83-84 


Onchocephalus 

nasutus  var.  cayennensis  243 

radiatus  243,  258 
Onchococephalus  234 
Oncocephalus  234 

longirostris  247 

notatus  243 

parvus  276 

porrectus  282 

radiatus  243,  259 

truncatus  243,  246 

vespertilio  252,  274 
Oncorhynchus 

sp.  423 

spp.  410 

Onychoteuthidae  427 
Onychoteuthis  425-426,  428 

sp.  421,  425 
Oostethus  184-185 

brachyurus  184-185 

lineatus  185 
Ophichthidae  57-67 
Ophichthinae  57-58,  63 
Ophichthus  62,  64 

(Coecilophis)  62 

erabo  57,  59,  65-66 

garretti  65 

kunaloa  57,  59,  61-62,  65-66 

pacifici  62 

polyophthalmus  57,  59,  64-66 

stypurus  63 

urolophus  62 
Ophichthys 

playfairii  64 

polyophthalmus  64 
Ophiodon  elongatus  417 
Ophisurus 

fasciatus  var.  semicinctus  64 

semicinctus  64 
Optonurus  atherodon  152 
Oreohelix  395 

elrodi  400 
Osmeridae  416-417 
Ostariophysi  497 

Pagophilus  429 
Palpigradida318 
Pandalopsis  427 
Pandalus  427 
Paracetonurus  135,  147,  149 

fragilis  138,  147-149 
Paralabrax  maculatofasciatus  220 
Parapercis  221 
Parasyngnathus  222-223 

elucens  184,  223 
Paratachys  455,  459 
Parhomaloptera  497,  500 


DEX 


fophrys  421 

vetulus  416 
giphaea  427 
ttella 

scurra  326 

striata  337 
telloida  324 

semirubida  328-329 

sm'ata  337 
yllinia  70 

Jicellaster  magister  megalabis  443-444,  449 
netopteryx  222 

taeniocephalus  184 
uenomonas  cooperae  57-58,  63,  66 
•anerodon  furcatus  417 
aromachrus  mocinno  473 
an,  409-410,  412,  414-415,  417-418,  428-429 

409,411,415 
412^13,  415^18,  428,  430 

phocoena  409,  411,  423-424 
ocoenidae  411 
xoenoides  412-413,  416-418 

dalli  411,  424-425 
dlophaga  4,  14-15,  42,  49 

/arcta  49 

rugosa  4,  37 

sp.  37,  39 

spp.  1 
"\\ophichthus  64 

xenodontus  47,  59,  64,  66 
xllopteryx  222-223 

/b/uttM.y  184,  223 
yseteridae  411 
ces  57,  135,  181,  287,  349 
oodonophis  magnified  63 
.tycephalidae  364 
lycephalus  indicus  364 
i/ynws  458 
ocoma  15 

urogrammus  monopterygius  415 
'aronectidae  409,  416-417 
'dlozonites  390,  400 

bermudensis  bermudensis  390 
'wees  428 
tf/es  instabilis  80 
licipes  polymerus  427 
meridius  459 
'ygonaceae  31,  33,  36 
ygyridae  401 
W/ia  4 

laponica  1,  4,  37 

fl/a  443-444 
409,  418,  421,  427 

notatus  409 

'idoryrhamphus  187 

lacksoni  187 

lacmaea  324 


547 


Proctocanthus  philadelphicus  45 
Progastromyzon  497 
Prokoenenia 
sp.  318 
Prosena  9-12 
siberita  3 

Prosenini  2,  5-7,  9-11,  15-16 
Prosenoides  11-12 
Prosopodasys  287 

asperrimus  287-290,  292 
Protomyzon  497,  500 
Pseudonezumia  135-136 
Pseudophallus  222 
elcapitanensis  184 
starksi  184 
Psocoptera  321 
Psydrini  455 
Ptilodexia  1-55 

abdominalis  4,  36-37 
o^i/w  3,  5,  10,  12,  14-15,  17-20,  24,  44 
/fo  group  12-13,  16,  18-23 

5,  9-10,  12-13,  15,  19,  45^6 
californica  1,  6-7,  9,  12,  15,  18-19,  29-31 
calif ornica  group  12-13,  29-36 
canescens  4-8,  12,  14,  16-17,  37-39,  42 
carolinensis  1,  4,  9,  11-12,  14-17,  19,  36-37,  42- 

43 

carolinensis  group  12-14,  36-41 
cerata  44 
confusa  44 

conjuncta  1,  3,  5-8,  12-14,  23-24,  48-49 
conjuncta  group  12-13,  23-29 
constrictans  26 

contristans  1,  5-7,  10,  12-13,  15-16,  18,  25-27 
dubia  44 
elevata  50 

flavotessellata  12-13,  15-17,  53-54 
fumipennis  48 

/ifl/om?  5-6,  8,  10,  12,  14,  17,  39-40 
harpasa  1,  4,  6,  9,  12-14,  18,  37-38,  41-44,  48,  50 
harpasa  group  12-13,  41-47 
hucketti  1,  39 

incerta  5-7,  12-14,  17-18,  49-51 
leucoptera  1,  41-42 

maculata  1,  4,  6-7,  10,  12-13,  15,  17,  51-53 
major  1,  11-13,  15,  18,  46-50 
major  group  12-13,  47-54 
mathesoni  5-6,  10,  12,  14,  17,  20-22 
minor  1,  37 

neotibialis  1,  36-37,  43 
obscura  9,  12-14,  19,  22-23,  41 
omissa  48 

pacifica  1,  5,  .7,  9-10,  12,  15,  19,  31-34 
planifrons  5-6,  8-10,  12-13,  15,  18,  24-25,  37,  42 
planifrons-contristans  5 
ponderosa  5,  8,  12-13,  17,  46-47 
prexaspes  4,  8-10,  12,  14,  17,  40-41 


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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol  4 


proximo  1,  50-51 

punctipennis  26 

rufipennis  1,  5-10,  12-14,  19,  42-45 

sabroskyi  1,  9,  12,  15,  18,  31,  34-36 

scutellata  48 

simulans  24 

spp.  3 

tibialis  11,  23,  41-42 

tincticornis  48 

westi  1,  5,  7,  10,  12-13,  15,  18,  27-29 
Ptilodexia-Mochlosoma  line  12 
Pulmonata  379 

Quietula  y-cauda  213,  219 

Raja  418 

[?]  Rana  piscatrix  251-253 
Rhacochilus  sp.  416 
Rhagium  lineatum  4 
Rhamnaceae  24,  45-46 
Rhamphinina  8,  11-12,  15-16,  49 

dubia  16,  48 

major  47-48 
Rhinotragides  69 
Rhinotraginae  69 
Rhinotragini  69-85 
Rhodopetala  324 
Rhynchiodexia  15-16,  44 

arida  45-48 

dubia  43-44 

flavotessellata  15,  53 

fumipennis  48 

levata  50 

mathesoni  20-21 

omissa  48 

ponderosa  46-47 

robusta  41-42 

tincticornis  47-48 
Rhynchodexia  15-16,  44,  50 

co/i/ura  1,  37-38,42^4 

conjuncta  23 

dubia  1 

elevata  1,  50 

fraterna  1,  48 

omissa  1,  48 

planifrons  24 

punctipennis  1,  25-26 

simulans  1,  23 

translucipennis  1,  43—44 
/«SSia?  423,  428 
Rosaceae  23 
Rubus  parviflorus  33 
Rudbeckia  sp.  24-25 
Rutiliini  11 

Salangidae  295,  297,  299,  302 
Salicaceae  20 


20 

Saperda  calcarata  4 
Sargassum  201,  216 
Scarabaeidae  14 
Scaritini  458 

Schismorhynchus  labialis  57-58,  62,  66 
Schultzidia  johnstonensis  58,  66 
Scomber  scombrus  423 
Scombridae  429 
Scorpaenidae  409,  416-417 
SCM/T/O  323-326,  337-338 

mesoleuca  326 

scurra  325-326 

stipulata  326 
Sebastes  418 

(Sebastomus)  capensis  complex  220 

(Sebastomus)  exsul-spinorbis  complex  220 

spp.  417-418 


salignus  24 

sp.  24,  27,  49 
Senostoma  9,  11,  16 
Serranidae  429 
Siphonostoma  198 
Siphostoma  198 

aciculare  207 

arc/Mm  193,  195 

auliscus  199 

bairdianum  211 

barbarae  211 

blainvilliana  190 

californiense  205 

californiensis  205 

carinatum  202 

exj/e  204 

griseolineatum  210 

leptorhynchum  210 

leptorhynchus  210 

sinaloae  201-202 
Sitalcina  sierra  316 
Solidago  3 

canadensis  22-23, 

juncea  37 

microcephala  41 

rugosa  42 

sp.  20,  24,  27,  33,  37,  45,  51 

trinervata  20,  25,  45 
Sphagebranchus 

flavicaudus  63 

vulturis  63 
Sphagemacrurus  135,  149 

frag  His  147 

trichiurus  171 
Sphagnum  466 
Spiraea  latifolia  23 
Spirinchus  421 
Spondias  72,  76 


42 


4DEX 


549 


qualus  418 
tanleya  pinnata  31 
tenopseutes  70,  80 
aeger  80 
sericinus  80 
indasalangidae  295-302 
mdasalanx  297,  299,  301-302 
microps  295-302 
praecox  295-302 
sp.  300-302 
mbolophorus  423 
nchiropus  349-350 

altivelis-group  350 

calauropomus  360 

phaeton-group  350 
ngnathidae  181-227 
ngnathiformes  181 
ngnathoides  222-223 

biaculeatus  184,  223 

tgnathus  181-182,  184,  192,  198-202,  204-205,  207- 
208,  213,  215,  218-220,  222 

abbotti  211-212 

acicularis  181,  190-192,  207 

acus  184-185,  198,  202,  207,  222 

agassizi  220 

arcta  195 

arctus  192,  195 

arundinaceus  211,  214 

auliscus  181,  184,  188,  199-202,  218,  220 
:  avus  181,  217-220 

bairdianus  185 

banner!  192 

barbarae  181,  210-212 

blainvilleanus  190 

blainvillei  190 

blainvillianus  190 

brevirostris  210-212,  215 

(Bryjt)  192,  199,  222 

(Bryx)  veleronis  192 

californiensis  181,  184-185,  199-201,  202-207,  211 
218,  220,  223 

californiensis  avus  219 

californiensis  californiensis  204,  208 

californiensis  griseolineatus  210 

californiensis  leptorhynchus  210 

carinatum  202 

carinatus  181,  184,  199-204,  207,  219-220 

can/iarwj-macrobrachium-/0//e///  lineage  219-220 

coccineus  192,  196-197 

darrosanus  192 

timidiatus  210-211 

iunckeri  192 

emeritus  181,  217-218,  220 

euchrous  181,  199-200,  206,  208-209,  211,  219-220 

euchrous  euchrous  201,  208-211,  220 

euchrous  intergrades  211 

euchrous  ollotropis  201,  208-211,  220 


emits  181,  199-201,  203-206,  219-220 
floridae  214 
folletti  203,  207,  219 
fuscus  198,  220 
griseolineatus  181,  205,  210 
griseolineatus  leptorhynchus  202 
hildebrandi  192 
hippocampus  185 
independencia  •  196-197 
insulae  181,  199-201,  209,  214-216,  219-220 
leptorhynchus  181,  184,  199-202,  205,  208-216 
218-220 

leptorhynchus-euchrous-insvl&e  lineage  220 
louisianae  214 

macrobrachium  181,  199-201,  203,  207-208,  219 
melanopleura  188 
(Microsyngnathus),  192,  195,  199 
norae  190 

pelagicus  184-185,  198,  218,  221 
phlegon  220 
randalli  192 

schlegeli  203-204,  214,  219 
scovelli  214 
spicifer  184-185 
spp.  220 

tweedlei  201-202 
typhle  212 
Syrictes  198 
exilis  204 

Tachinidae  1-2,  5,  9,  11,  36 
Tachyina  455,  458-459 
Tamaricaceae  49 
Tamarix  sp.  49 
Tarletonbeania  423 
Tectura  324-325 

rosacea  328 
Tetraclita  529 
Thaleichthys  pacificus  416 
Theragra  chalcogramma  416,  423 
Thoracica  527 
Thunnus  alalunga  204 

albacares  187 

thynnus  187 
Tomopterus  70 

exilis  69-71 

staphylinus  70 

vespoides  70-71 
Topobea  308 

micrantha  303,  307-308 
Trachonurus  135 
Trechini  455 
Trichodon  423,  427 
Trigonia  floribunda  77 
Trogloneta  paradoxa  318 

t//va  428 


550 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES,  Vol. 


Umbelliferae  45 
Umbellularia  californica  33 
Urocyclinae  399 
Urophori  184,  222-223 
Usofila  (Telema)  sp.  318 

Ventrifossa  135,  149,  151-153,  157,  163 
atherodon  151-152 
(Atherodus)  152 
barbiger  157 
ctenomelas  152 
diver  gens  152 
fasciata  153 
garmani  152 
lucifer  152-153 
(Lucigadella)  152 
(Lucigadus)  135,  149,  151-153 
(Lucigadus)  nigromaculata  138,  153 
(Lucigadus)  sp.  cf.  ori  153 
macropogon  152-153 
misakia  153 
mucocephalus  152-153 


nigromaculata  138,  153-156,  177 

nigromarginata  152-153 

ori  153 

petersonii  152 

(Sokodara)  135,  151-153 

(Ventrifossa)  135,  151-152 
Verbenaceae  24 
Verbesina  encelioides  26 
Verma  flavicauda  63 

Watasenia  424 

. 

Xanthoxylum  74 
Xenaploactis  287-293 

flrtopta  287,  289-291 

asperrima  289-290 

cautes  287-288,  291-292 

Zalieutes  234 

Zalophus  410,  412,  414-415,  417-419,  422,  428-429 
californianus  409,  411,  419,  421-422 


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